九州大学総合研究博物館研究報告第17号
Bulletin of the Kyushu University Museum
No.17, 2020, pp.1-108

Taxonomic revision of the ant genus Myrmecina in Southeast Asia
(Hymenoptera: Formicidae)
1)2)

Hirofumi OKIDO , Kazuo OGATA & Shingo HOSOISHI
1)

2), *

Kyushu Environmental Evaluation Association, 1-10-1 Matsukadai, Higashi-ku, Fukuoka, 813-0004 Japan
2)

Institute of Tropical Agriculture, Kyushu University, 744 Motooka, Nishi-ku, Fukuoka, 819-0395 Japan＊

Corresponding author: E-mail: hosoishi@agr.kyushu-u.ac.jp

Abstract: The ant genus Myrmecina occurring in Southeast Asia is systematically revised. Fifty-three species of this genus
are recognized from these areas. The following 40 species are described as new to science: M. arcuata, M. asiatica, M. aspera,
M. asthena, M. boltoni, M. breviata, M. celebebsis, M. dasynota, M. dechai, M. dolichothrix, M. elegans, M. glabra, M. gopa,
M. gracilis, M. grandis, M. gymnocephala, M. inflata, M. insulana, M. inthanonensis, M. itoi, M. lambirensis, M. lombokensis,
M. longiseta, M. macrops, M. mahuana, M. maryatiae, M. monticola, M. muluensis, M. nigra, M. nomurai, M. padangensis,
M. parallela, M. poringensis, M. sabahna, M. spinosa, M. sulawesiana, M. sundanica, M. tridentata, M. vieti and M. yamanei.
A key to the species based on the workers is provided. Of all species examined, 16 species are classified into four species
complexes, though the remainings are not assigned to any species groups.
Key Word: Myrmicinae, Crematogastrini, taxonomy, new species

Introduction
The ant genus Myrmecina of the subfamily Myrmicinae was established by Curtis (1829) based on Myrmecina latreillii Curtis.
This genus is assigned to the tribe Crematogastrini Forel, 1893 (Ward et al. 2015; Bolton, 2020). This genus consists of 56 valid
species at present (Bolton, 2020) and is distributed in all the geographical regions except for the Ethiopian region. Members of thisgenus are medium-sized, characterized by a pair of longitudinal carina on the ventrolateral margin of the head, and cylindrical petiole
without an anterior peduncle.
Of the 56 nominal species, 28 were described from Asia (Bolton, 2020). This figure shows that Myrmecina species are the richest
in Asia. The previous studies, however, were fragmental and treated with only local faunas based on limited materials i.e., Japan(Ogata & Terayama, 1992; Terayama & Kihara, 1994), China and Taiwan (Lin & Wu, 1998; Zhou, 2001; Huang et al. 2008; Zhou etal. 2008), India (Tiwari, 1994), Thailand (Jaitrong et al. 2019), Singapore (Wong & Guénard, 2016), Borneo (Wheeler, 1919), Sumatra(Forel, 1913; Satria & Yamane, 2019).
Biology of the genus Myrmecina has been studied in only a few species. Masuko (1994) reported the predatory habits of M.
nipponica and M. flava, and showed that the ants are specialized predators on oribatid mites. He concluded that the peculiarly elongate head of the larvae is associated with a feeding habit in which they insert the head into the hard oribatid body to consume thecontents. The association of oribatid mites with Myrmecina was also observed in tropical Asia (Ito & Takaku, 1994; Aoki & Ito, 1997)
where they show symbiosis. The colony size of the ants has been recorded in the limited number of species. Wilson (1959) noted 50
adults in a colony of M. transversa. Ohkawara et al. (1993) counted 27.4±SD 16.1 adults per colony from 108 samples of M.
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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

nipponica. Ito (1996) recorded 66±SD 24 workers and 8±SD 8.4 ergatogynes per colony from 41 samples of Myrmecina species ofJava. Ergatogynes are found in other species. Emery (1916), Ohkawara et al. (1993) and Ito (1996) reported that M. graminicola, M.
nipponica and undescribed species of Java have ergatogynes.
Members of the genus Myrmecina live in rotten wood (Wilson, 1959), soil and litter, or under stones (Ogata & Terayama, 1992).
The ants are relatively rare, but have a potential value as a bioindicator especially in tropical Asia, because they occur well conserved
forests. The taxonomy of the genus is, however, far from complete. Up to the present, no comprehensive reviews have been made,which causes practical difficulties in identification of the group. It is necessary to study its taxonomy for the inventory of this area.
The purpose of the present study is to establish a classification system of the genus Myrmecina in Southeast Asia. The study area
includes the entire Oriental region, and the Austro-Malayan subregion of the Australian region.

Material and methods‘Southeast Asia’ in the present paper refers to the Oriental region and Australian region excluding Australia, Papua New Guinea,
the Solomon Islands, Vanuatu, Fiji and the islands in the Pacific Ocean.
Specimens examinedType specimens were examined and/or deposited in the collectioons below. Other specimen depositories are also listed (Someof abbreviations of these institutions follow those of Brandão, 2000).
AMNH		 American Museum of Natural History, New York, U. S. A.
BMNH		 The Natural History Museum, London, United Kingdom.
BZM			 Bogor Zoological Museum, Bogor, Indonesia.
EUMJ			 Entomological Laboratory, Ehime University, Matsuyama, Japan.
HNHM		 Hungarian Natural History Museum, Budapest, Hungary.
IEBR			 Institute of Ecology and Biological Resources, Hanoi, Vietnam.
KUEC			 Entomological Laboratory and Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan.
KUFA			 Faculty of Agriculture, Kagawa University, Miki, Japan.
KUFF			 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok, Thailand.
KUIC			 Department of Earth and Environmental Sciences, Faculty of Science, Kagoshima University, Kagoshima, Japan.
LKCNHM Lee Kong Chian Natural History Museum, Singapore.
MCSN			 Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy.
MCZC			 Museum of Comparative Zoology, Harvard University, Massachusetts, U. S. A.
MHNG		 Museum d’Histoire Naturelle, Geneva, Switzerland.
MNHA		 Museum of Nature and Human Activities, Hyogo, Japan.
MTPC			 Mamoru Terayama, Private Collection, Tokyo, Japan.
MZB			 Bogor Zoological Museum, Bogor, Indonesia.
NHMW		 Naturhistorisches Museum Wien, Vienna, Austria.
NIAES			 National Institute for Agro-Environmental Sciences, Tsukuba, Japan.
NSMT			 National Science Museum, Tokyo, Japan.
NMNS			 National Museum of Natural Science, Taichung, Taiwan.
NTU			 Department of Plant Pathology and Entomology, National Taiwan University, Taipei, Taiwan.
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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

NZSI			 National Zoological Collection, Zoological Survey of India, Calcutta, India.
OMNH		 Osaka Museum of Natural History, Osaka, Japan.
RSC				 Rijal Satria Collection, Padang, West Sumatra, Indonesia.
SBSHKU		Insect Biodiversity and Biogeography Laboratory, School of Biological Sciences, The University of Hong Kong, Hong
Kong SAR.
SKYC			 Seiki Yamane Collection, Kagoshima, Japan.
TARI			 Taiwan Agricultural Research Institute, Taichung, Taiwan.
THNHM		 Thailand Natural History Museum, Technopolis, Khlong Luang, Pathum Thani Thailand.
UMS			 Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Kota Kinabalu, Malaysia.
USNM			 United States National Museum of Natural History, Washington, D. C., U. S. A.
The specimens used in this study were collected mainly from the following Asian countries:
Brunei, China, Indonesia (Sumatra, Java, Sebesi, Sulawesi), Malaysia, Myanmar, Philippines, Thailand, Vietnam. The material collected by a colony sampling method was shown with colony code in various ways.
Observation of morphological features
Observations were made under stereoscopic microscopes (Nikon, SMZ-2T; Olympus, SD-STB3) and a scanning electron microscope (SEM: JEOL, JSM-5600LV) at voltage 15 kV. In the later case, some specimens were sputter-coated with gold for 12 min (4min × 3 times), but the others not.
Measurements and indicesMeasurements, indices and their abbreviations follow those of Bolton (2000).
Total Length (TL): The total length of the ant from the mandibular apex to the gastral apex; when measured in profile the sum of
mandible length + HL + ML + lengths of waist segments (petiole and postpetiole) + length of gaster.
Head Length (HL): The length of the head capsule excluding the mandibles, measured in full-face view along mid-dorsal longitudinal axis from the mid-point of the anterior clypeal margin to the mid-point of the occipital margin. In species where one or bothof these margins are concave, the measurement is taken from the mid-point of a transverse line that spans the apices of the projecting
portions.
Head Width (HW): The maximum width of the head in full-face view, measured behind the eyes.
Cephalic Index (CI): HW*100 divided by HL.
Scape Length (SL): The maximum straight-line length of the antennal scape, excluding the basal constriction or neck close to thecondylar bulb.
Scape Index (SI): SL*100 divided by HW.
Pronotal Width (PW): The maximum width of the pronotum in dorsal view.
Mesosoma Length (ML) (=Weber’s length, alitrunk length): The diagonal length of the mesosoma in profile from the point at
which the pronotum meets the cervical shield to the posterior basal angle of the metapleuron.
Measurements were made by means of a stereoscopic microscope (SMZ-2T) with an ocular graticule and expressed in millimeters (mm). The scale bars in the digital photographs by SEM were expressed in micrometers (µm).

―3―Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Morphology and terminology
Note on the terminology concerning the castes in Myrmecina
In the present study, taxonomy of the genus Myrmecina is based on the morphological characters of the workers. But in somespecies there are individuals morphologically intermediate between the queen and worker, and wingless as workers. It is sometimes
difficult to distinguish workers from them. Therefore, it is appropriate to state about the wingless females and its terms.
These wingless females have been called “ergatogynes” (Ohkawara et al., 1993). Peeters (1991) suggested that there are two
distinct categories in the term ergatogynes: ergatoid queens and intercastes. The ergatoid queens are a persistently wingless reproductive caste which has derived from normal queens, while the intercastes are morphologically continuous series of reproductive females.
Here, these categories are not recognized, and “ergatogyne” is used only when the wingless females can be morphologically distinguished from workers. Because morphological difference between ergatoid queen and intercaste is not clear, and their ontogenetic
process is not known.
Morphological differences between worker and ergatogyne are observed mainly in total body length, head width, eye length,
shape, width and sutures of mesosoma, and width of gaster. Among 53 species examined in this study, ergatogynes are found in six
species: M. gopa sp. nov., M. grandis sp. nov., M. guangxiensis Zhou, M. magnificens Wong & Guénard, M. raviwonghei Jaitrong etal., M. sundanica sp. nov. (see the remarks under each species). In addition, Forel (1915) and Emery (1916) reported the occurrenceof ergatogyne in the European species, M. graminicola (Latreille, 1802).
Morphology of worker
Head
The head is usually rectangular with rounded occipital corners (Figs. 3d), but in some species, the corners are projected posteriorly (Fig. 42e). The ratio of the length to width of the head varies among species. The occipital margin is straight or concave at themedian portion; the occipital carina is present. The ventrolateral margin is delineated by a sharp longitudinal carina on each side,which arises close to inner-ventral mandibular base, running along the length of the head immediately below the eye and ascending
to the occipital surface posteriorly. This character is unique to the genus Myrmecina. The lobate area partly covering an antennal
socket is called the frontal lobe, and the longitudinal ridge following the frontal lobe is called the frontal carina. In many species of
the genus, the frontal lobe is more or less distinct, but the frontal carina is short, weak and almost indistinguishable from the surface
sculptures. The median portion of the clypeus projects with or without small processes on the anterior margin (Fig. 1a); the numberof the small processes ranges from 1 to 4; the lateral portions are usually simple, but in the M. vieti complex with sharp ridges of shieldwalls in front of the antennal insertions. Eyes are variable in size and the number of ommatidia, obscure in some some species complexes but distinct in others. The eye is usually located at the apical one third between the mandibular insertion and the occipitalcorner of the head in profile. Because of the variable size of the eye, the shortest distance between the base of the mandible and theanterior margin of the eye, or the malar space, varies among species.
The antenna is 12-segmented, forming a club of apical 3 segments. The basal portion of the antennal scape bears a more or lessdeveloped flange (Fig. 1a), which encircles the area above the basal condylar bulb. This structure, the antennal flange (Moffett,1986), is seen in the genera Acanthomyrmex, Myrmecina, Perissomyrmex and Pristomyrmex. The antennal flange is extremely welldeveloped in M. flava Terayama, 1985, completely covering the condylar bulb. The third segment of the antenna lacks the girdle ofhairs. Taylor (1980) noticed this character in M. cacabau (Mann, 1921) and thought it as a distinctive character of this species. This
character was observed in all the species examined, and thus this is a diagnostic character of the genus.
The mandible is robust and strongly curved downwards. As the distal portion of each mandible in anterior view is suddenly
broadened, this portion is distinguished easily from the proximal portion forming an arm. The shape of the mandible in the genus is―4―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

occipital cornerantennal flange

basal tooth

small teeth
third toothmalar space
preapical tooth
apical toothab

median process of median portionof anterior clypeal margin
denticle of dorsolateral
portion of pronotum
pronotal lobe

eumetanotal spine
propodeal
spiracledenticle of anterior
ventrolateral portionof pronotum

propodeal
lobe
subpetiolar
processcright angled corner ofposterior ventrolateral
portion of pronotum

mesoepisternal projection

Fig. 1. Morphological terms. a, head of M. spinosa sp. nov. in full-face view; b, masticatory marginof mandible of M. maryatiae sp. nov. in anterior view; c, mesosoma and petiole of M. spinosa sp.
Fig.in 1profile.
Morphological terms. a, head of M. spinosa sp. nov. in fullnov.

face view; b, masticatory margin of mandible of M. maryatiae sp.
anterior
view;c, andmesosomapetioleof M.margin
spinosa
sp. bent at mid-length, but
subtriangular, havingnov.
three inmargins:
the outer-,basalmasticatory andone. The
masticatory
is usuallynov.
inprofile.
sometimes straight, and armed with a strong apical, the preapical and the third teeth distinctly. The preapical tooth is incomplete andfused with apical one, without isolated tip (Fig. 1b). These three teeth are followed by small teeth varying in number and a blunt basal
tooth (Fig. 1b). The basal tooth is sometimes absent. The dorsal surface of the labrum bears projected structure in the genera Acanthomyrmex, Myrmecina, Perissomyrmex and Pristomyrmex. In the genus Myrmecina, paired denticles usually occur on the anteromedian portion, but sometimes the projections are fused with each other at the base. The number of segments of the maxillary and labial
palpi, or palp formula, is consistently 4 and 3, respectively.
MesosomaIn this study, the term ‘mesosoma’ is used for calling the second tagma including pro-, meso-, metathorax and propodeum. The
term is synonymus with ‘trunk’ and ‘alitrunk’ of myrmecology. The mesosoma of the genus Myrmecina is flattened to strongly
convex on the dorsal margin in profile. The anterior portion of the pronotum is usually marginated and broadened (Fig. 3a). The
anterior dorsolateral and ventrolateral portions are armed with or without denticles. Those denticles are well developed in M. spinosa,―5―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

and its allied species (Fig. 1c). Even when the denticles are not formed, the anterior ventrolateral portion is distinct and angulate. The
posterior ventrolateral portion extends and forms a right angled corner. The upper most area of the free posterior margin of the pronotum forms a rounded lobe covering the mesothoracic spiracle, which is homologous with pronotal lobe in other hymenopterans.
The mesonotum is completely fused with the pronotum, lacking the promesonotal suture. The mesopleuron is fused with themesonotum. Its border is probably originated from the area just posterior to the pronotal lobe. The anterior ventrolateral portion
extends to form a projection partly covering the forecoxa, here termed the mesoepisternal projection. The posterior ventrolateral
portion of the pronotum and the mesoepisternal projection form a cleavage. The oblique furrow on the mesoepisternum is absent,
which is present in the male and female. The line was errorously interpreted as the suture dividing the anepisternum and the katepisternum in Bolton (1994).
In some cases of myrmicines there is a groove between the mesonotum and the propodeum. This represents a vestige of themetanotum, and thus called the metanotal groove. In the worker of Myrmecina, however, the metanotal groove is completely lost.
The most species of Myrmecina have a pair of spines in front of the propodeal spines. The metanotum of the female has a pair ofprocesses in some species. Comparing the worker with the conspecific female as shown in M. spinosa sp. nov. (Figs. 2a, 2b), thespines in front of the propodeal ones in the worker seem to be originated in the metanotal area. Thus, the spines in front of theabFig. 2. Eumetanotal spines of M. spinosa sp. nov. in oblique view. a, worker; b,queen.

Fig. 2 Eumetanotal spines― 6 of― M. spinosa sp. nov. in
oblique view. a, worker; b, queen.

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

propodeal ones are “metanotal spines”. But the term “metanotal spines” was used incorrectly in the past literature of myrmecology(e.g., Bingham, 1903), here I newly introduce the term “eumetanotal spines” to this structure (Fig. 1c).
The propodeal spines are usually triangular to elongate, but only angulately projected (not spine-like) in M. breviata sp. nov. and
M. dasynota sp. nov. The propodeal lobe (Fig. 1c) is a small process developed on the propodeal declivity. It varies among species
from a low ridge to distinct projection. The propodeal spiracles are variable in size and its position among species, usually situatednear the base of the propodeal spines; distance between the posterior margin of the spiracle and that of the propodeum is usually longer
than the diameter of the spiracle.
The legs are relatively short. The middle and hind legs lack the tibial spurs.
Petiole, postpetiole and gaster
The petiole is the second abdominal segment morphologically. The tergum and the sternum are rigidly united; the former has
spiracles anteroventrally; the sternum is reduced to form a simple longitudinal plate, often with a small anterior projection termed
subpetiolar process. In almost all the myrmicine genera, the segment is petiolate, including an anterior peduncle and a posterior node.
But in the genus Myrmecina, the petiole is sessile, somewhat cylindrical in overall shape, so that the peduncle and the node are indistinguishable; the anterior portion is truncated forming an oblique face, which is marginated by a curved ridge in profile; the highest
portion, the dorsal crest, is located at the midlength of the petilole. The shape of the petiole and the proportion of the height and length
are useful characters in distinguishing species. The height is defined as the minimum length taken from the dorsal crest to ventral edgeof the petiole in profile. The length is defined as the minimum length taken from the anterior most point of the curved ridge to theposterior margin of the petiole in profile.
The postpetiole is subrectangular in dorsal view, with a transverse anterior margin and subparallel sides. The lateral margins are
sometimes convergent. The proportion of the width to length is different by species. The anterior portion of the postpetiolar sternum
is sharply marginated, sometimes forming a distinct projection in profile.
The gaster is usually oval in dorsal view. In dorsal view, the base of the first gastral tergum is simple in many species, but insome species the portion is concave behind the postpetiole and forming distinct anterolateral corners. The first gastral tergum is large,
occupying about two thirds of the dorsal area of the gaster. The sting is present but small and weakly sclerotized. The morphologyof the sting apparatus was studied by Kugler (1978, 1979). No stinging behavior has been observed (Maschwitz, 1964; Masuko,1994).
Surface
The sculpture on the body surface in this genus is one of conspicuous characters distinguishing it from other genera easily. In
the majority of species, the head and mesosoma have longitudinal or irregular rugae distinctly. In many cases, patterns of the sculptureof them are more or less different among species. Thus, this sculpturing character is used for recognizing species in the following key
to the species, although the sculpture is variable in some species.
The dorsal portion of the head has longitudinal or irregular rugae with or without punctures. On the ventrolateral portion of thehead, some species have sculptured surface and others have no sculpture. The scuplture of this portion is sometimes used for recognizing species. The ventral portion of the head has transverse and curved rugae in almost all speices.
The dorsal and the lateral portions of the mesosoma have usually longitudinal or irregular rugae with or without punctures,
sometimes transverse and curved rugae or diverging rugae anteriorly. The sculpture of the head and mesosoma is variable between
species, sometimes the sculpture is not observed and the surface of the head and mesosoma is smooth and shining in some species.
The petiole and postpetiole usually have longitudinal rugae on the dorsal and lateral portions. The first gastral segment is usuallysmooth and shining, sometimes shallow rugae or punctures are observed in a few species.
―7―Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

The body surface including the mandible, antenna and legs has erect pilosity. The pilosity is simple but variable in density andlength among species. In particular, the ratio of its length on the pronotum to length of propodeal spine is sometimes useful for recognizing species.
Body color is usually black, but the color of the legs, the antenna and the posterior portion of the gaster is often reddish or yellowish brown. The color of the whole body is sometimes yellowish brown, or slightly darker in M. dasynota sp. nov.

Taxonomy
Genus Myrmecina Curtis
Myrmecina Curtis, 1829: 265. Type species: Myrmecina latreillii Curtis (junior synonym of Myrmecina graminicola) by monotypy.
Archaeomyrmex Mann, 1921: 448. Type species: Archaeomyrmex cacabau Mann, by original designation [synonymy by Brown,
1971: 1.].
Diagnosis of workers
Occipital corners rounded in profile but sometimes projected posteirorly. Ventrolateral margin delineated by a sharp longitudinalcarina on each side, the carina arising close to inner-ventral mandibular base, running along length of head immediately below eye
and ascending to occipital surface posteriorly. Frontal lobe present. Mandible robust and strongly down-curved with basal arm;
anterior clypeal margin and basal margin of mandible forming a space when the mandibles are closed in full-face view; masticatory
margin with a strong apical, low and blunt preapical and third teeth followed by 4-6 small teeth and a blunt basal tooth. Apical toothalways acute and distinct; preapical tooth fused with apical one, sometimes indistinct; third tooth well isolated from preapical one;basal tooth sometimes reduced in size and indistinguishable from median small teeth. Clypeus more or less projected anteriorly, withor without 1-4 processes on median portion of anterior margin. Labrum with a pair of denticles on median anterodorsal portion, eachof which are distinct and well separated. Antenna 12-segmented, with apical 3 segments forming a club; third segment lacking thegirdle of hairs; base of scape forming flange more or less covering condylar bulb. Palp formula 4, 3.
Pronotum with or without denticles on dorsolateral or ventrolateral portions. Mesoepisternal projection covering a part of forecoxa. Eumetanotal spines usually present but often small to absent.
Petiole cylindrical, without an anterior peduncle.
First gastral tergum large, occupying about two thirds of dorsal area of gaster, with or without sculptures dorsally, anterior margin
sometimes concave.
Note[Spelling]
There was a nomenclatural confusion on the type species. When Curtis (1829) established the genus Myrmecina, he originallydescribed M. latreillii based on a male. In his catalogue, Bolton (1995) listed up this as a type species of Myrmecina. But he also used
the same species as “M. latreillei” at the section of the species list. Latter spelling has been cited by Mayr (1855), Nylander (1856),
Emery (1895), Wheeler (1911), Rigato (1999) and others. This is incorrect spelling according to the Article 32 and 33 of ICZN(ICZN, 1999).
[Synonymy]
Bolton (1995) showed that Mayr (1855) synonymized latreillei [sic] under graminicola. But the truth is that Mayr treated―8―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

latreillei as a senior synonym of graminicola. Indeed Emery (1895) used the species name latreillei when he described the subspecies
americana. Mayr’s treatment was wrong, because he based it on the incorrect authorship of graminicola as Foerster (1850). The
name, Myrmecina graminicola, has been used as valid since Emery (1898) (see also Donisthorpe, 1915; Emery, 1922).
Checklist of the species of Southeast Asia
In the present study, 53 species are treated and listed here arranged by alphabetical order. In the list below, 40 species of them
are new to science, and their specific names are followed by the boldface, “sp. nov.”. Sixteen species of them could be categorized
to 4 species complexes. These complexes are indicated by the abbreviations in the brackets following each specific name as follows:[gra], gracilis complex; [mac], macrops complex; [spi], spinosa complex; [vie], vieti complex. The other species are not assigned toany of these complexes. The distribution of species is indicated following each specific name.
1.		 andalas Satria & Yamane [spi]: INDONESIA: Sumatra
2.		 arcuata sp. nov. PHILIPPINES: Luzon.
3.		asiatica sp. nov. CHINA: Yunnan Prov.; THAILAND: Nakhon Ratchasima Prov., Chanthaburi Prov., Phetchaburi Prov., Pattani
Prov.; MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra, Rakata I.
4.		 aspera sp. nov. MALAYSIA: The Malay Pen.
5.		 asthena sp. nov. CHINA: Yunnan Prov.; VIETNAM: Lao Cai Prov.; MALAYSIA: The Malay Pen., Borneo (Sabah).
6.		 bandarensis Forel INDONESIA: Sumatra.
7.		 boltoni sp. nov. INDONESIA: Sulawesi.
8.		 breviata sp. nov. [mac] PHILIPPINES: Luzon.
9.		 butteli Forel INDONESIA: Sumatra.
10. celebensis sp. nov. INDONESIA: Sulawesi.
11. curvispina Zhou et al. [gra] CHINA; VIETNAM: Vinh Phuc Prov., Ninh Binh Prov.
12. dasynota sp. nov. MALAYSIA: Borneo (Sabah).
13. dechai sp. nov. THAILAND: Phetchaburi Prov.
14. dolichothrix sp. nov. PHILIPPINES: Mindanao.
15. elegans sp. nov. MALAYSIA: Borneo (Sabah).
16. glabra sp. nov. MALAYSIA: The Malay Pen.
17. gopa sp. nov. INDONESIA: Java.
18. gracilis sp. nov. [gra] MALAYSIA: Borneo (Sabah, Sarawak); INDONESIA: Borneo (Kalimantan).
19. grandis sp. nov. MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); BRUNEI: Tasek Merimbum; INDONESIA: Sumatra.
20. guangxiensis Zhou [vie] CHINA; VIETNAM: Ninh Binh Prov.
21. gymnocephala sp. nov. [mac] PHILIPPINES: Mindoro; INDONESIA: Seram.
22. inflata sp. nov. [spi] THAILAND: Phang Nga Prov.
23. insulana sp. nov. MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); INDONESIA: Sumatra, Borneo (Kalimantan), Java.
24. inthanonensis sp. nov. THAILAND: Chiang Mai Prov.
25. itoi sp. nov. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra.
26. lambirensis sp. nov. MALAYSIA: Borneo (Sarawak).
27. lombokensis sp. nov. INDONESIA: Lombok.
28. longiseta sp. nov. INDONESIA: Sumatra.
29. macrops sp. nov. [mac] MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra, Java.
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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

30. magnificens Wong & Guénard [spi] SINGAPORE; MALAYSIA: Tioman Is., The Malay Pen.; INDONESIA: Sumatra.
31. mahuana sp. nov. [spi] MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra.
32.	
maryatiae sp. nov. THAILAND: Phang Nga Prov.; MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); INDONESIA:
Borneo (Kalimantan).
33. monticola sp. nov. MALAYSIA: The Malay Pen.
34. muluensis sp. nov. MALAYSIA: Borneo (Sarawak).
35. nesaea Wheeler INDONESIA: Sebesi I. (near Krakatau Is.)36. nigra sp. nov. VIETNAM: Lao Cai Prov.
37. nitidiuscula Satria & Yamane [mac] INDONESIA: Sumatra.
38. nomurai sp. nov. [vie] VIETNAM: Cao Bang Prov., Vinh Phuc Prov.
39. padangensis sp. nov. INDONESIA: Sumatra.
40. parallela sp. nov. INDONESIA: Sumatra.
41. poringensis sp. nov. MALAYSIA: Borneo (Sabah).
42. raviwonghei Jaitrong et al. [gra] THAILAND: Chiang Mai Prov., Necornratchasima Prov.
43. sabahna sp. nov. MALAYSIA: Borneo (Sabah).
44. semipolita Forel INDONESIA: Java.
45.	
spinosa sp. nov. [spi] MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); BRUNEI: Tasek Merimbun; INDONESIA:
Borneo (Kalimantan), Java.
46. striata Emery MYANMAR: Tenasserim.
47. sulawesiana sp. nov. INDONESIA: Sulawesi, Seram.
48. sulcata Emery INDONESIA: Sulawesi, Java.
49. sundanica sp. nov. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Java, Bali.
50. tridentata sp. nov. [spi] MALAYSIA: Borneo (Sabah, Sarawak).
51. undulata Emery INDONESIA: Sumatra, Laut I. (SE of Borneo).
52. vieti sp. nov. [vie] VIETNAM: Ha Tai Prov.
53. yamanei sp. nov. MALAYSIA: The Malay Pen.
Myrmecina gracilis complex
The M. gracilis complex is characterized by the small eyes (< 0.12 mm), the flattened mesosoma, long and posteriorly directed
propodeal spine, and first gastral tergum that is concave at the anterior margin in dorsal view and sculptured on the dorsum. The
species complex comprises M. curvispuna Zhout et al., 2008 M. gracilis sp. nov. and M. raviwonghei Jaitrong et al., 2019, and is
distributed in N. Vietnam, Thailand and Borneo.
The propodeal spine extends to the vertical posteriormost limit of the propodeum, and is at the almost same level of the flattened
mesosoma. The apex of the spine is acute and directed posteriorly, though some variations occur with species. The long, acute andhorizontal spine is apomorphic, but also observed in some species such as M. spinosa sp. nov. and M. tridentata sp. nov. The concave
anterior margin of the gaster forms rounded lateral corner in each side. This state is apomorphic, but again observed in several species
in other species grous such as M. aspera sp. nov., M. elegans sp. nov., M. spinosa sp. nov. and M. tridentata sp. nov. The species ofthis complex tend to be small in size.
Myrmecina macrops complex
The M. macrops complex is characterized by having the large eyes, the broadened head, and the smooth and shining, or foveolate― 10 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

surface of the head. The species complex includes M. breviata sp. nov., M. gymnocephala sp. nov., M. macrops sp. nov. and M. nitidiuscula Satria & Yamane, 2019. The species complex is distributed in the Philippines, Sumatra, Malay Peninsula, Borneo, Java.
All these characters seemed to be derived states but each of them can be observed independently in different species complexesof the genus. M. maryatiae sp. nov. would be close to the complex in having the similar shape of the head, but the shape of the pronotum and the surface sculpture are different from those in the complex.
Myrmecina spinosa complex
The M. spinosa complex is characterized by the backwardly projected occipital corners of the head (Fig. 42e). The species
complex includes M. andalas Stria & Yamane, 2019, M. inflata sp. nov., M. magnificens Wong & Guénard, 2016, M. mahuana sp.
nov., M. spinosa sp. nov. and M. tridentata sp. nov. The species complex is distributed in Thailand, S. Malay Peninsula, Borneo,
Sumatra and Java.
This unique shape of the head is an autapomorphy of the complex and somewhat similar to that of Gnamptogenys species of the
Ectatomminae. Species of this complex are relatively larger than other species. Among 5 species of the complex, M. spinosa sp. nov.
and M. tridentata sp. nov. share the distinct projections on the mesosoma including anterior dorsolateral and anterior ventrolateral
denticles on the pronotum, the eumetanotal spines and the propodeal spines (Figs. 42b, 46b), the shape of the masticatory margin of
the mandible, and the concave anterior margin of the first gastral tergum.
Myrmecina vieti complex
The M. vieti complex is well defined as having a sharp ridge of the shield wall on the lateral portion of the clypeus in front of theantennal insertions. The species complex includes M. guangxiensis Zhou, 2001, M. nomurai sp. nov. and M. vieti sp. nov. Distributionof the species complex is restricted to E. Asia.
The shield wall is unique and represents a derived state among the genus Myrmecina. In the tribe Crematogasterini, Acanthomyrmex has an incomplete ridge originated from the frontal lobe but low and indistinct in front of the antennal insertion. Pristomyrmex and Perissomyrmex have a distinct ridge surrounding the frontal area of the antennal insertion, but the anterior margin of theclypeus hides under this ridge. The shield wall in the M. vieti complex and its ridge have the same property judging from their position
and the connection to the frontal lobe. At least the structure in the M. vieti complex would be homologous among the members,
because the anterior margin of the clypeus does not hide but is visible. Another character demarcating the M. vieti complex is theshape of the propodeal lobe (Figs. 20b, 36b, 47b). The lobe is reduced and low in this complex, while is more or less distinct in others.
The same state is also observed in several species including M asthena sp. nov., M. breviata sp. nov., M. dasynota sp. nov., M. dechai
sp. nov., etc. This would be a secondary reduction. From these characters discussed above, the M. vieti complex would be a monophyletic group having these synapomorphies.
Key to species based on workers
The key presented below treats 48 species. The following 5 species are not examined and omitted from the key: butteli Forel,1913 from Sumatra, nesaea Wheeler, 1924 from Sebesi I., semipolita Forel, 1905 from Java, striata Emery, 1889 from Myanmar,
undulata Emery, 1900 from Sumatra.
1.		 Lateral portions of clypeus raised into a sharp ridge of shield wall in front of the antennal insertions������������2 (M. vieti complex)-		 Lateral portions of clypeus not raised into a sharp ridge of shield wall in front of the antennal insertions������������������������������������42.		 Petiole longer than high; lateral surface smooth; ventral margin concave in profile (Fig. 47b)����������������������������������� vieti sp. nov.
-		 Petiole as long as high; lateral surface rugose; ventral margin not concave in profile (Fig. 36b)���������������������������������������������������3― 11 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

3.		Body color reddish brown. Anterior portion of postpetiole forming more or less distinct lateral expansion, broadest anteriorly
and much broader than petiole. Mesosoma with more or less flattened dorsal margin in profile; dorsal surface with longitudinaland parallel rugae����������������������������������������������������������������������������������������������������������������������������������������������������nomurai sp. nov.
-		Body color black. Anterior portion of postpetiole not expanded laterally, slightly broader than petiole. Mesosoma with convex
dorsal margin in profile; dorsal surface with rugae diverging anteriorly������������������������������������������������������������ guangxiensis Zhou
4.		 Occipital corners of head projected posteriorly (Fig. 42e)���������������������������������������������������������������������������5 (M. spinosa complex)-		 Occipital corners of head not projected posteriorly (Fig. 7b)��������������������������������������������������������������������������������������������������������10
5.		 First gastral tergum not concave in dorsal view (Fig. 31a)��������������������������������������������������������������������������������������������������������������6-		 First gastral tergum concave in dorsal view (Figs. 22a, 46a)�����������������������������������������������������������������������������������������������������������7
6.		Mesosoma in lateral view with dorsal outline evenly convex. Mesosoma with irregular longitudinal rugulae. Propodeal spine
pointing dorsally����������������������������������������������������������������������������������������������������������������������������������������andalas Satria & Yamane-		Mesosoma in lateral view with dorsl outline highly convex. Mesosoma with irregular rugulae. Propodeal spine pointing posteriorly���������������������������������������������������������������������������������������������������������������������������������������������������������������������� mahuana sp. nov.
7.		Masticatory margin of mandible straight (Fig. 22d)�������������������������������������������������������������������������������������������������� inflata sp. nov.
-		Masticatory margin of mandible bent at midlength (Fig. 1b)����������������������������������������������������������������������������������������������������������8
8.		Propodeal spine elevated, pointing dorsally; propodeal declivity almost vertical with posterodorsal angle about 90˚		����������������������������������������������������������������������������������������������������������������������������������������������������������� magnificens Wong & Guénard
-		Propodeal spine extending backward and over vertical posteriormost limit of propodeum in profile; propodeal declivity steepwith posterodorsal angle < 90˚ (Figs. 42b, 46b)�������������������������������������������������������������������������������������������������������������������������������99.		Head and mesosoma with straight and longitudinal rugae on dorsal surface (Figs. 42a, 42c). Postpetiole distinctly marginate,sometimes with projected lateral lobe on both sides in dorsal view; anterior dorsal margin angulately raised in profile		���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� spinosa sp. nov.
–		Head and mesosoma with wavy or irregular rugae on dorsal surface (Figs. 46a, 46c), sometimes the rugae on mesosoma transverse. Postpetiole not marginate on both sides in dorsal view; anterior dorsal margin not angulately raised, almost horizontal inprofile�������������������������������������������������������������������������������������������������������������������������������������������������������������������� tridentata sp. nov.
10.	Head smooth and shining (Fig. 21c), or sparsely foveate (Fig. 29c) on dorsal surface�����������������������������������������������������������������11-		 Head largely sculptured with rugae and sometimes dense punctures on dorsal surface (Figs. 3c, 5c, 6c, 40c)����������������������������15
11.	Head longer than broad (Fig. 16c)������������������������������������������������������������������������������������������������������������������������������glabra sp. nov.
-		Head as long as broad (Fig. 21c)�������������������������������������������������������������������������������������������������������������� 12 (M. macrops complex)
12.	Propodeal spine only angulately projected, not spine-like (Fig. 9b)����������������������������������������������������������������������� breviata sp. nov.
-		Propodeal spine well developed, triangular to elongate (Fig. 29b)������������������������������������������������������������������������������������������������13
13.	Head sparsely foveate on dorsal surface�����������������������������������������������������������������������������������������������������������������macrops sp. nov.
-		Head smooth and shining, or mostly smooth and shining except for frontal carinae and few rugulae in front of eye������������������14
14.	Mesosoma with straight rugulae in dorsal view; median longitudinal portion raised, forming arched surface		�����������������������������������������������������������������������������������������������������������������������������������������������������������������������gymnocephala sp. nov.
-		Mesosoma with wavy rugulae in dorsal view������������������������������������������������������������������������������������ nitidiuscula Satria & Yamane
15.	Anterior margin of first gastral tergum more or less concave in dorsal view��������������������������������������������������������������������������������16-		Anterior margin of first gastral tergum not concave in dorsal view�����������������������������������������������������������������������������������������������20
16.	First gastral tergum sculptured�������������������������������������������������������������������������������������������������������������������17 (M. gracilis complex)-		First gastral tergum not sculptured�������������������������������������������������������������������������������������������������������������������������������������������������1917.	First gastral sternum not sculptured�������������������������������������������������������������������������������������������������������������� curvispina Zhout et al.
-		First gastral sternum sculptured�����������������������������������������������������������������������������������������������������������������������������������������������������18― 12 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

18.	First gastral sternum with ridge at anterior median portion in ventral view. Anterior clypeal margin without single medianprocess. Head longer than broad����������������������������������������������������������������������������������������������������������������������������� gracilis sp. nov.
-		First gastral sternum without ridge at anterior median portion in ventral view. Anterior clypeal margin with single medianprocess. Head as long as broad���������������������������������������������������������������������������������������������������������������raviwonghei Jaitrong et al.
19.	Propodeal spine broad and triangular, 1.5 times as long as broad at base or shorter, just reaching vertical posteriormost limit ofpropodeum in profile. Pronotum with rugae on dorsal surface. Eye comprising 15 ommtidia or more����������������� aspera sp. nov.
-		Propodeal spine elongate, much longer than broad at base, extending over vertical posteriormost limit of propodeum in profile.
Pronotum smooth and shining on dorsal surface. Eye comprising 10 ommatidia or less��������������������������������������� elegans sp. nov.
20.	Antennal scape long, extending beyond occipital corner in full-face view�����������������������������������������������������������������������������������21-		Antennal scape short, not or just reaching occipital corner in full-face view��������������������������������������������������������������������������������31
21.	Eumetanotal spine absent���������������������������������������������������������������������������������������������������������������������������������������������������������������22-		Eumetanotal spine present��������������������������������������������������������������������������������������������������������������������������������������������������������������23
22.	Anterior clypeal margin without processes�������������������������������������������������������������������������������������������������������dolichothrix sp. nov.
-		Anterior clypeal margin with three processes�������������������������������������������������������������������������������������������������������������� gopa sp. nov.
23.	Ventral portion of petiole simple, without subpetiolar process nor median longitudinal ridge (Figs. 28b, 33b)���������������������������24-		Ventral portion of petiole with subpetiolar process or median longitudinal ridge (Figs. 1c, 44b)�������������������������������������������������28
24.	Anterior clypeal margin with median paired processes�����������������������������������������������������������������������������������������������������������������25
		 Anterior clypeal margin with single median process or without processes�����������������������������������������������������������������������������������26
25.	Head with straight and thick rugae on dorsal surface (Fig. 8c). Forecoxa with rugae����������������������������������������������boltoni sp. nov.
-		Head with slightly waved and thin rugae, and punctures on dorsal surface (Fig. 33c). Forecoxa smooth and shining		������������������������������������������������������������������������������������������������������������������������������������������������������������������������������ monticola sp. nov.
26.	Propodeal spine long, extending over vertical posteriormost limit of propodeum in profile���������������������������������parallela sp. nov.
-		Propodeal spine short, not or just reaching vertical posteriormost limit of propodeum in profile (Figs. 28b, 33b)����������������������27
27.	Pronotal pilosity sparse and extremely long, almost three times as long as propodeal spine (Fig. 28b). Dorsal portion of propodeal
lobe not raised, simple. Ventrolateral portion of head sculptured (Figs. 3d, 23d). Eye much shorter than malar space		��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������longiseta sp. nov.
-		Pronotal pilosity dense and short, twice as long as propodeal spine or shorter (Fig. 33b). Dorsal portion of propodeal lobe raised,
forming rounded corner. Ventrolateral portion of head smooth and shining (Fig. 7b). Eye as long as malar space		�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������bandarensis Forel
28.	Ventrolateral portion of head sculptured (Figs. 3d, 23d)����������������������������������������������������������������������������������������������������������������29-		Ventrolateral portion of head smooth and shining (Fig. 7b)�����������������������������������������������������������������������������������������������������������30
29.	Anterior clypeal margin with single median process. Mesosoma with straight and parallel rugae on dorsal surface (Fig. 24a). Eyecomprising 20 ommatidia or more����������������������������������������������������������������������������������������������������������������� inthanonensis sp. nov.
-		Anterior clypeal margin with three processes. Mesosoma with wavy and diverging rugae on dorsal surface (Fig. 35a). Eye comprising 15 ommatidia or less����������������������������������������������������������������������������������������������������������������������������������������nigra sp. nov.
30.	Eye small, comprising 15 ommatidia or less; the diameter much shorter than half of malar space. Head longer than broad (CI<90)(Fig. 34c)�������������������������������������������������������������������������������������������������������������������������������������������������������������� muluensis sp. nov.
-		Eye large, comprising 20 ommatidia or more; the diameter longer than half of malar space. Head as long as broad (CI≥98)		�����������������������������������������������������������������������������������������������������������������������������������������������������������������������sulcata Emery (p. 118)
31.	Pilosity extremely dense (Fig. 12b); number of hairs on petiole more than 20�����������������������������������������������������dasynota sp. nov.
-		Pilosity sparse (Figs. 13b, 39b); number of hairs on petiole less than 15��������������������������������������������������������������������������������������32
32.	Mesosoma with transverse rugae on dorsal surface�����������������������������������������������������������������������������������������������������������������������33― 13 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI-		Mesosoma with longitudinal or irregular rugae on dorsal surface�������������������������������������������������������������������������������������������������34
33.	Propodeal spine long, pointed backward��������������������������������������������������������������������������������������������������������������������dechai sp. nov.
-		Propodeal spine short, pointed upward��������������������������������������������������������������������������������������������������������������poringensis sp. nov.
34.	Dorsal median portion of mesosoma and lateral portion of pronotum not sculptured���������������������������������������lambirensis sp. nov.
-		Dorsal median portion of mesosoma and lateral portion of pronotum sculptured�������������������������������������������������������������������������35
35.	First gastral sternum sculptured���������������������������������������������������������������������������������������������������������������������������celebensis sp. nov.
-		First gastral sternum not sculptured�����������������������������������������������������������������������������������������������������������������������������������������������36
36.	First gastral tergum sculptured�����������������������������������������������������������������������������������������������������������������������������sundanica sp. nov.
-		First gastral tergum smooth and shining����������������������������������������������������������������������������������������������������������������������������������������37
37.	Eye distinctly longer than half of malar space�������������������������������������������������������������������������������������������������������������������������������38-		Eye as long as or shorter than half of malar space�������������������������������������������������������������������������������������������������������������������������40
38.	Head broad posteriorly in full-face view (Fig. 37c). Petiole distinctly longer than high in profile. Rugae on dorsal surface of headrunning parallel and straight (Figs. 8c, 37c)����������������������������������������������������������������������������������������������������� padangensis sp. nov.
-		Head with lateral margins nearly parallel in full-face view, anterior width almost same as posterior one (Figs. 25c, 32c). Petiole
almost as long as high in profile. Rugae on dorsal surface of head running wavy and irregularly (Figs. 25c, 32c)���������������������39
39.	Masticatory margin of mandible straight without basal tooth�����������������������������������������������������������������������������������������itoi sp. nov.
-		Masticatory margin of mandible bent at the midlength with basal tooth (Fig. 1b)���������������������������������������������� maryatiae sp. nov.
40.	Masticatory margin of mandible straight (Fig. 19d)������������������������������������������������������������������������������������������������ grandis sp. nov.
-		Masticatory margin of mandible bent at midlength (as in Fig. 1b)������������������������������������������������������������������������������������������������41
41.	Rugae on ventrolateral portion of head transverse (Fig. 23d)�������������������������������������������������������������������������������� insulana sp. nov.
-		Rugae on ventrolateral portion of head longitudinal or absent (Figs. 3d, 7b)��������������������������������������������������������������������������������42
42.	Antennal scape not reaching occipital corner by scape width in full-face view (SI≤77)����������������������������������������yamanei sp. nov.
-		Antennal scape just reaching occipital corner in full-face view (SI≥80)���������������������������������������������������������������������������������������43
43.	Anterior clypeal margin with median paired processes������������������������������������������������������������������������������������������sabahna sp. nov.
-		Anterior clypeal margin with single median process or without processes�����������������������������������������������������������������������������������44
44.	Ventrolateral portion of head with punctures, without longitudinal rugae�������������������������������������������������������������������������������������45-		Ventrolateral portion of head without punctures, or with both punctures and longitudinal rugae (Fig. 7b)����������������������������������46
45.	Eye small and circular, comprising about 6 ommatidia; the diameter much shorter than half of malar space. Head longer thanbroad�������������������������������������������������������������������������������������������������������������������������������������������������������������������������asthena sp. nov.
-		Eye medium-sized and oval, comprising about 14 ommatidia; the diameter as long as half of malar space. Head as long as broad�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� lombokensis sp. nov.
46.	Eye small and circular, comprising about 9 ommatidia; its diameter much shorter than half of malar space���������asiatica sp. nov.
-		Eye medium-sized and oval, comprising 15 ommatidia or more; its diameter as long as half of malar spacesulawesiana sp. nov.
Species accounts

Myrmecina andalas Satria & Yamane
Myrmecina andalas Satria & Yamane, 2019: 185: Holotype worker, INDONESIA: Sumatra, Aceh, Leuser Ecosystem, Putri Betung,
Gunung Kemiri, N 3°49’521’’, E 97°31’198’’, alt. ca. 950–1200 m, 19.ix.2012 (R. Satria) (SEMUT20180326A) (MZB). Paratypes.
One worker (SEMUT20180731A), same data as holotype (RSC); 3 workers (SKYU-SI-FOR001–003), Leuser Ecosystem, Gunung― 14 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Kemiri, alt. 1200 m, good forest, 20.ix.2012 (Yamane & Syaukani) (SKYC); 1 ergatoid queen (SEMUT20180731B), same data as
holotype (RSC). [one SKYC paratype worker examined].
Remarks. M. andalas belongs to the M. spinosa complex. Among those species, M. andalas can be distinguished from the remainsby having the elevated propodeal spine and simple anterior margin of the first gastral tergum.
Distribution. INDONESIA: Sumatra.

Myrmecina arcuata sp. nov.
(Fig. 3)
Holotype worker. TL 2.62, HL 0.66, HW 0.59, CI 90, SL 0.56, SI 95, PW 0.432, ML 0.72.
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth unclear. Dorsal surface of clypeus notslightly concave; median portion of anterior margin projected with three median processes that are very small; lateral portion simple,
lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles very small. Frontal carinae
virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and convex with maximum diameter 0.10 mm, with 6
ommatidia in the longest row; malar space slightly twice as long as diameter of eye or shorter in profile; distance between occipitalmargin and posterior margin of eye four times as long as diameter of eye. Antennal scape long, extending beyond posterolateralcorner of head; antennal flange developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior portion marginate; anterior ventrolateral
portion angulate. Furrow between pronotum and mesoepisternal projection narrow. Eumetanotal spine absent. Propodeal spine
elongate, 1.5 times as long as broad at base, extending over vertical posteriormost limit of propodeum in profile. Propodeal lobe low.
Propodeal spiracle large, situated near posterior margin of propodeum, distance between posterior margin of spiracle and posterior
margin of propodeum slightly longer than diameter of spiracle. Petiole long, longer than high in profile, and longer than broad in
dorsal view; dorsal crest absent; subpetiolar process absent. Postpetiole slightly broader than petiole in dorsal view; anterodorsal
outline slightly convex in profile; ventral outline projected with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae on dorsum which are distinct and longitudinal; ventrolateral portion with deep straight rugae. Clypeus
smooth and shining. Mesosoma with deep straight rugae longitudinally. Forecoxa smooth and shining. Petiole and postpetiole witha few distinct rugae. First gastral segment smooth and shining. Head with relatively sparse and long pilosity on dorsum. Mesosoma
with relatively sparse and long pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral
surface. Postpetiole with 1-2 hairs on ventral surface. Body black or dark reddish brown, forecoxae reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, PHILIPPINES: Luzon, Lagunas, Mt. Banahaw, 24. xi. 1995, 700 m, (I. Löbl) (BMNH).
Remarks. M. arcuata does not belong to any species complex. But in terms of the relative length of the antennal scape, the shape of― 15 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

a

b

d
c
Fig. 3. Myrmecina arcuata sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view ofhead; d, ventrolateral portion of head.

the mesosoma without the eumetanotal spine, and the shape of the first gastral tergum, M. arcuata is similar to M. modesta Mann andM. subarmata Mann. The similarity of those species does not always reflect the relationships, because the characters mentioned above
tend to change independently. Among those species, M. arcuata can be distinguished from the remains by having the elongate propodeal spine.

― 16 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Distribution. PHILIPPINES: Luzon.

Myrmecina asiatica sp. nov.
(Fig. 4)
Holotype worker. TL 2.53, HL 0.59, HW 0.54, CI 92, SL 0.50, SI 91, PW 0.38, ML 0.64.
Paratype workers. TL 2.51-2.68, HL 0.59-0.61, HW 0.54-0.56, CI 91-95, SL 0.50, SI 89-91, PW 0.37-0.39, ML 0.66-0.70 (5abcFig. 4. Myrmecina asiatica sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

― 17 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

measured).
Worker. Head subrectangular, slightly longer than broad, sometimes as long as broad in full-face view; median portion of occipitalmargin concave; occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small
tooth or sixth tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus notconcave; median portion of anterior margin feebly projected with or without a median process; lateral portion simple, lacking sharp
ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles, which are usually well separated.
Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes extremely small, varying in size with maximum
diameter 0.06 mm and 7-11 ommatidia, circular, and moderately convex; malar space usually three times as long as diameter of eye
or longer, but variable; distance between occipital margin and posterior margin of eye distinctly five times as long as diameter of eye
or longer, but variable. Antennal scape short, just reaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion incompletely marginate;
anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine
present but small, sometimes unclear. Propodeal spine triangular, variable in length, usually extending beyond, sometimes just reaching vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior marginof propodeum, usually apart from margin by its diameter. Petiole short, as long as or slightly longer than high in profile, and slightly
longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process variously developed from weakly
raised median longitudinal ridge to distinct projection. Postpetiole slightly broader than petiole in dorsal view; dorsal outline flattenedor slightly convex in profile; ventral outline not or slightly projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view.
Head with slightly waved or straight rugae, sometimes also punctured; ventrolateral portion usually with longitudinal rugae and/or punctures, sometimes smooth and shining. Clypeus smooth and shining. Mesosoma with deep and longitudinal rugae with orwithout punctures. Forecoxa smooth and shining. Petiole and postpetiole usually with longitudinal rugae, sometimes with transverse
or irregular rugae, or almost smooth and shining. First gastral segment smooth and shining. Head with dense and short pilosity on
dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum usually shorter than propodeal spine. Petiole without
hairs on ventral surface. Postpetiole usually with 1-2 hairs on ventral surface. Head and mesosoma black, forecoxae reddish brownto yellowish brown, petiole, postpetiole and gaster black to reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, CHINA: Dahaoping, Gaoligongshan Mts., Tengchong Xian, Yunnan Prov., 10. x. 1996 (S. Nomura) (KUEC).
Paratypes. 7 workers with same data as holotype (NSMT, KUEC, KUIC).
Other material examined. MALAYSIA: Malaya, Selangor, Gombak (C. Betts); Neg. Sembilan, Pasoh For. Res. (M. Brendell, K.
Jackson & L. Ficken) (BMNH); Sabah, Kinabalu N. P., Poring Hot Springs, East Ridge (C. Brühl); Sabah, Kinabalu (Burckhardt &Löbl) (BMNH); Sabah, Crocker Range, KK-Tambunan, 1600 m (Löbl & Burckhardt) (BMNH); Sabah, Crocker Range, 1650 m (Löbl& Burckhardt) (BMNH); Sabah, KK-Tambunan, 1550 m-1650 m (Löbl & Burckhardt) (BMNH); Sabah, Crocker Range N. P., Gg.
Emas Highland Res. (no collector’s name). THAILAND: Sai Khao, Pattani Prov. (H. Okido); Kaeng Krachan NP. (Löbl & Burckhardt) (BMNH); Khao Soi Dao W. S., Chanthaburi Prov. (H. Okido); Kao Yai, Nakornratchasrima Prov. (Misspelling; Nakhon Ratchasima) (W. Jaitrong; Löbl & Burckhardt) (BMNH); Doi Chiang Dao, 500-600 m alt., nr. Chiang Mai (H. Okido); Doi Inthanon(Löbl & Burckhardt) (BMNH). INDONESIA: N. Sumatra, Brastagi, 1500 m (Rougemont); Sitiung, W. Sumatra (F. Ito); Sumatra,
Padang (no collector’s name) (BMNH); Rakata I., 150 m alt., Krakatau Is., S 06°09´xE 105°27´ (H. Simbolon); Rakata I., 50 m alt.,
Krakatau Is., S 06°09´xE 105°27´ (K. Ogata).
― 18 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Remarks. M. asiatica does not belong to any species complex. But in terms of the body size, the masticatory margin of the mandible,
relative length of the antennal scape, the shape of first gastral tergum, and the body sculpture, the species is similar to M. sulawesiana
sp. nov. The similarity of those species does not always reflect the relationships, because the characters mentioned above tend tochange independently. Although differences between these two species are subtle, M asiatica can be distinguished from the latter byhaving small and circular eye comprising about 9 ommatidia. Some morphological characters of the species are variable, and thespecies is widely distributed in the Indo-Chinese and Indo-Malayan subregions. In the light of these facts, it may be possible that the
species would be split into more than one species.
The species inhabits in soil and litter.
Distribution. CHINA: Yunnan Prov.; THAILAND: Nakhon Ratchasima Prov., Chanthaburi Prov., Phetchaburi Prov., Pattani Prov.;
MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra, Rakata I. (Krakatau Is.).

Myrmecina aspera sp. nov.
(Fig. 5)
Holotype worker. TL 3.14, HL 0.80, HW 0.77, CI 96, SL 0.77, SI 100, PW 0.54, ML 0.85.
Paratype worker. TL 3.06, HL 0.79, HW 0.75, CI 95, SL 0.72, SI 96, PW 0.51, ML 0.82 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin distinctly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible straight; apical tooth strong, third tooth robust, followedby unclear 4 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portion of anterior margin projected,
with or without three processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface oflabrum with paired denticles varies in size and shape, sometimes fused with each other at base. Frontal carinae virtually absent,
indistinguishable from rugae on dorsum of head. Eyes large and convex with maximum diameter 0.14 mm, with 7 ommatidia in thelongest row; malar space slightly longer than diameter of eye in profile; distance between occipital margin and posterior margin of eye
three times as long as diameter of eye or shorter. Antennal scape long, extending beyond posterolateral corner of head; antennal flange
fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum with denticles on dorsolateral portion, but small; anterior
portion marginate; anterior ventrolateral portion projected directing downward. Furrow between pronotum and mesoepisternal projection narrow. Eumetanotal spine present but small. Propodeal spine broad and triangular, 1.5 times as long as broad at base or
shorter, just reaching vertical posterior most limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle small, situatednear the base of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole short, slightly longer than high in profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process absent. Postpetiole as broad as petiole, and expanded posteriorly in dorsal view; dorsal outline
flattened; ventral outline slightly projected rectangularly in profile.
Anterior margin of gaster concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with deep wavy rugae; ventrolateral portion with transverse, or vertical rugae posteriorly. Clypeus smooth and shining.
Mesosoma with deep wavy rugae. Forecoxa almost smooth and shining. Petiole and postpetiole with a few rugae. First gastralsegment smooth and shining. Head with relatively sparse and long pilosity on dorsum. Mesosoma with sparse and long pilosity on
dorsum, hairs of pronotum much longer than propodeal spine. Dorsal pilosity of petiole as long as that of mesosoma. Petiole without― 19 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 5. Myrmecina aspera sp. nov. a, dorsal view of mesosoma,
petiole and postpetiole; b, lateral view of mesosoma, petiole andpostpetiole; c, full-face view of head.

hairs on ventral surface. Postpetiole with 4 hairs on ventral surface. Head and mesosoma black, petiole, postpetiole and gaster reddish
brown, mandibles, antennae and legs yellowish brown to light reddish brown.
Holotype worker, MALAYSIA: Genting Highlands, 28. ix. 73 (B. Bolton) (BMNH). Paratype. 1 worker with same data as holotype(BMNH).
Other material examined. MALAYSIA: Fraser Hill, Pahang Prov., 1180 m (Schillhammer) (NHMW).
― 20 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Remarks. M. aspera does not belong to any species complex. But in terms of the body size and the smooth and shining first gastral
tergum with the concave anterior margin, M. aspera is similar to M. elegans sp. nov. The similarity of those species does not always
reflect the relationships, because the characters mentioned above tend to change independently. Among those species, M. aspera canbe distinguished from the remains by having the broad and triangular propodeal spine, the sculptured dorsum of the pronotum, andthe eye comprising 15 ommatidia or more.
Distribution. MALAYSIA: The Malay Pen.

Myrmecina asthena sp. nov.
(Fig. 6)
Holotype worker. TL 3.07, HL 0.72, HW 0.62, CI 87, SL 0.54, SI 87, PW 0.43, ML 0.77.
Paratype workers. TL 2.86-2.98, HL 0.64-0.69, HW 0.56-0.61, CI 86-88, SL 0.51-0.56, SI 87-92, PW 0.38-0.42, ML 0.74-0.78 (5measured).
Worker. Head subrectangular, longer than broad distinctly in full-face view; median portion of occipital margin concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin feebly projected, with or without three processes; lateral portion simple, lacking sharp ridge in front of antennal
insertions. Anterior dorsal surface of labrum with paired small denticles, which are well separated. Frontal carinae absent. Eyes
extremely small circularly and not convex, varying in size with maximum diameter 0.05-0.06 mm and 5-8 ommatidia; malar space
three times as long as diameter of eye or longer in profile; distance between occipital margin and posterior margin of eye distinctly
five times as long as diameter of eye or longer. Antennal scape short, just reaching posterolateral corner of head; antennal flange
developed.
Dorsal outline of mesosoma convex to flattened in profile. Pronotum without denticles; anterior portion not marginate completely in dorsal view; anterior ventrolateral portion not angulate. Mesoepisternal projection reduced; cleavage between pronotum
and mesoepisternal projection broad. Eumetanotal spine present but small. Propodeal spine triangular, variable in length, usuallyextending over vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near baseof propodeal spine, apart from margin by its diameter. Petiole long, longer than high in profile, and longer than broad in dorsal view;dorsal crest located at midlength in profile; subpetiolar process with acute anterior apex, or median longitudinal ridge at ventral
portion present. Postpetiole slightly broader than petiole in dorsal view; dorsal outline flattened in profile; ventral outline clearly
projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view.
Head densely punctured sometimes with unclear longitudinal rugae; ventrolateral portion usually punctured, rarely unclearly.
Clypeus smooth and shining. Mesosoma with thin and straight rugae weakly. Forecoxa smooth and shining. Petiole and postpetiole
with longitudinal rugae. First gastral segment smooth and shining. Head with dense and short pilosity on dorsum. Mesosoma with
dense and short pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 4 hairs on ventral surface. Body black to reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, CHINA: Dabei, 2440 m alt., Gaoligongshan Mts., Tengchong Xian, Yunnan Prov., 11. x. 1996 (S. Nomura)― 21 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 6. Myrmecina asthena sp. nov. a, dorsal view of mesosoma, petiole and postpetiole;b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

(KUEC).
Paratypes. 11 workers with same data as holotype. CHINA: 1 worker, Dahaoping Gaoligongshan Mts., Tengchong Xian, Yunnan
Prov., 10. x. 1996 (S. Nomura); 1 worker, Huanxipo, Tengchong Xian, Yunnan Prov., 14. x. 1996 (S. Nomura) (KUEC, NSMT).
Other material examined. VIETNAM: Sin Chai, 1620 m alt., Sapa, Lao Cai Prov. (B. T. Viet); Thac Bac, 1820 m alt., Sapa, Lao Cai
Prov. (B. T. Viet) (KUEC). MALAYSIA: Sabah, Crocker Range NP., Gg. Emas Highland Res., 1500-1700 m (no collector’s name);
Malaya, Gombak (B. Bolton) (BMNH).
― 22 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Remarks. M. asthena does not belong to any species complex. But in terms of the head sculpture, relative length of the antennal
scape, process of the clypeus, and shape and sculpture of the first gastral segment, M. asthena is similar to M. lombokensis sp. nov.
The similarity of those species does not always reflect the relationships, because the characters mentioned above tend to change independently. Among those species, M asthena can be distinguished from the remains by having the small and circular eye comprising
about 6 ommatidia.
Distribution. CHINA: Yunnan Prov.; VIETNAM: Lao Cai Prov.; MALAYSIA: The Malay Pen., Borneo (Sabah).

Myrmecina bandarensis Forel(Fig. 7)Myrmecina bandarensis Forel, 1913: 72. Syntype workers and queen, INDONESIA: Bandar Baroe, Sumatra, No. 598 (V. Buttel)(MHNG) [syntypes examined].
Worker. TL 2.48-2.60, HL 0.59-0.61, HW 0.60-0.62, CI 101-103, SL 0.51-0.56, SI 85-90, PW 0.37-0.38, ML 0.64-0.69 (3measured).
Worker. Head subrectangular, shorter than broad in full-face view; median portion of occipital margin slightly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median
portion of anterior margin slightly projected with a median process; lateral portion simple, lacking sharp ridge in front of antennal
insertions. Anterior dorsal surface of labrum with paired denticles, fused with each other at base. Frontal carinae virtually absent,
indistinguishable from rugae on dorsum of head. Eyes extremely large and convex, and located at anterior portion near the mid pointof head in profile, varying in size with maximum diameter 0.14-0.16 mm and 7-8 ommatidia; malar space as long as diameter of eye
in profile; distance between occipital margin and posterior margin of eye 1.5 times as long as diameter of eye. Antennal scape long,extending beyond posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion not marginate; anterior
ventrolateral portion more or less projected; anterior dorsolateral portion rounded. Furrow between pronotum and mesoepisternal
projection broad. Eumetanotal spine present but small. Propodeal spine triangular, longer than broad at base, just reaching verticalposteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine,distance between posterior margin of spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole short, aslong as high in profile, and slightly longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar processabsent.

Postpetiole as broad as petiole in dorsal view; dorsal outline flattened in profile; ventral outline slightly projected

rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae which are thick and longitudinal; ventrolateral portion smooth and shining. Clypeus smooth andshining. Mesosoma with straight rugae which are thick and longitudinal. Forecoxa smooth and shining. Petiole and postpetiole witha few rugae. First gastral segment smooth and shining. Head and mesosoma with relatively sparse and long pilosity on dorsum, hairsof pronotum as long as propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 4 hairs on ventral surface. Body
brown to reddish brown, mandibles, antennae, and legs yellowish brown.
― 23 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 7. Myrmecina bandarensis Forel. a, dorsal view of mesosoma, petiole and postpetiole;b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Remarks. M. bandarensis does not belong to any species complex. But in terms of the large eye and the head shape, the species
is similar to the species of the macrops complex and M. maryatiae sp. nov. The similarity of those species does not always reflect therelationships, because the characters mentioned above tend to change independently. M bandarensis can be distinguished from thelatters by having the sculptured head and the long antennal scape.
Distribution. INDONESIA: Sumatra.
― 24 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Myrmecina boltoni sp. nov.
(Fig. 8)
Holotype worker. TL 3.87, HL 0.91, HW 0.88, CI 96, SL 0.87, SI 99, PW 0.59, ML 1.09.
Paratype workers. TL 3.51-4.46, HL 0.84-1.08, HW 0.82-1.04, CI 93-100, SL 0.80-0.91, SI 88-100, PW 0.53-0.67, ML 1.04-1.22 (6measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin flattened or slightly concave;abcFig. 8. Myrmecina boltoni sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

― 25 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth
tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus concave;median portion of anterior margin slightly projected with two or four processes; lateral portion simple, lacking sharp ridge in front ofantennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and distinctly convex, varying in size with maximum diameter 0.11-0.16 mm and 5-10 ommatidia; malar space longer than diameter of eye in profile; distance between occipital margin andposterior margin of eye twice as long as diameter of eye or longer. Antennal scape long, extending beyond posterolateral corner ofhead; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum with small denticles but unclearly; anterior portion not marginate; anterior ventrolateral portion usually angulate, sometimes not angulate. Furrow between pronotum and mesoepisternal projection
narrow, sometimes relatively broad. Eumetanotal spine long. Propodeal spine triangular, variable in length, not or just reaching vertical posteriormost limit of propodeum in profile; as long as broad at base or longer. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum longer thandiameter of spiracle. Petiole long, longer than high in profile, and slightly longer than broad in dorsal view; dorsal crest located at
midlength in profile; subpetiolar process and median longitudinal ridge usually absent. Postpetiole slightly broader than petiole indorsal view; dorsal outline flattened or slightly convex in profile; ventral outline slightly projected with two tops.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae distinctly; ventrolateral portion with also deep and straight rugae longitudinally but frequntly smoothand shining. Clypeus smooth and shining. Mesosoma with thick rugae longitudinally. Forecoxa with transverse rugae. Petiole andpostpetiole with longitudinal rugae. First gastral segment smooth and shining. Head with sparse and long pilosity on dorsum. Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum longer than propodeal spine. Petiole with or without 1-4 hairs on
ventral surface. Postpetiole with 2-8 hairs on ventral surface. Head, mesosoma, forecoxae, petiole and postpetiole black, gasterreddish brown, mandibles, antennae and legs yellowish brown to brown.
Holotype worker, INDONESIA: Sulawesi, Utara, G. Muajat, 1780 m, 16. ix. 1985 (no collector’s name) (BMNH).
Paratypes. 4 workers and 1 male with same data as holotype; 3 workers with same data as holotype but 25. i. 1985; 2 workers withsame data as holotype but Summint area, ca 1780 m, 24. v. 1985 (BMNH).
Other material examined. INDONESIA: Sulawesi, Utara, Dumoga-Bone N. P., 12?. ix. 1985 (no collector’s name) (BMNH).
Remarks. M. boltoni does not belong to any species complex. But in terms of the simple petiole without the subpetiolar process nor
median longitudinal ridge, relative length of the antennal scape, and the mesosoma with the eumetanotal spine, M. boltoni is similar
to M. monticola sp. nov. The similarity of those species does not always reflect the relationships, because the characters mentioned
above tend to change independently. Among those species, M boltoni can be distinguished from the remains by having the anterior
clypeal margin with median paired processes and the sculptured forecoxa.
Distribution. INDONESIA: Sulawesi.

Myrmecina breviata sp. nov.
(Fig. 9)― 26 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Holotype worker. TL 2.60, HL 0.65, HW 0.61, CI 94, SL 0.61, SI 100, PW 0.42, ML 0.70.
Paratype worker. TL 2.59, HL 0.66, HW 0.64, CI 98, SL 0.59, SI 93, PW 0.42, ML 0.70 (1 measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin flattened; occipital corners
widely rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a stout basal tooth bearing blunt apex; small teeth unclear. Dorsal
surface of clypeus slightly concave; median portion of anterior margin weakly projected with a median process but the process frequently unclear; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum withpaired small denticles distinctly, closed to each other. Frontal carinae very short, frequently unclear, running to about mid point levelabcFig. 9. Myrmecina breviata sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

― 27 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

of eyes. Eyes extremely large and convex, varying in size with maximum diameter 0.15-0.16 mm and 7-8 ommatidia; malar space aslong as diameter of eye in profile. Antennal scape long, extending beyond posterolateral corner of head distinctly; antennal flange
weakly developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior ventrolateral portion not angulate.
Eumetanotal spine absent. Propodeal spine only angulately projected, not spine-like. Propodeal lobe feebly raised. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum
slightly longer than diameter of spiracle. Petiole long, longer than high in profile, and longer than broad in dorsal view; small but
distinct dorsal crest located almost at midlength in profile; subpetiolar process absent or unclear; ventral outline flattened or slightly
convex. Postpetiole broader than petiole in dorsal view; anterior side corners rounded; dorsal outline slightly convex or flattened in
profile; ventral outline projected rectangularly.
Anterior margin of gaster not concave in dorsal view.
Head smooth and shining; ventrolateral portion smooth and shining. Mandibles smooth and shining, frequently yellow spots on
dorsal surface. Clypeus smooth and shining. Mesosoma largely smooth and shining, lateral portion with a few longitudinal rugae on
dorsum. Petiole with shallow rugae. Postpetiole smooth and shining on dorsum. Head with sparse pilosity on dorsum. Mesosoma
with sparse pilosity on dorsum. Pilosity of pronotum slightly shorter than maximum diameter of eye. Pilosity of petiole and postpetiole as long as that of mesosoma. Petiole without hairs on ventral surface. Postpetiole with about 4 hairs on ventral surface. Head,mesosoma, petiole and postpetiole reddish brown, gaster slightly darker than head and mesosoma.
Holotype worker, PHILIPPINES: Luzon, Lagunas, Mt. Makiling, 21. xi. 1995 (I. Löbl) (BMNH).
Paratype. 1 worker with same data as holotype (BMNH).
Other material examined. PHILIPPINES: Luzon, Lagunas, Mt. Banahaw above Kinabuhayan, 600-700 m, trail to Crystalino (J.
Kodada & B. Rigová) (NHMW).
Remarks. M. breviata forms the macrops complex comprising M. breviata, M. gymnocephala sp. nov. and M. macrops sp. nov. This
species is different from the related species in having the angulately projected propodeal spine.
Distribution. PHILIPPINES: Luzon.

Myrmecina butteli Forel
Myrmecina butteli Forel, 1913: 71. Syntype worker, INDONESIA: Tandjong Slamat, Sumatra (Buttel-Reepen?) (depository unknown)[not seen].
Diagnosis of worker. The following description is based on Forel (1913). Mandible smooth and shining. Masticatory margin ofmandible with 6-7 teeth. Anterior margin of clypeus slightly concave. Mid point of head length as long as broad. Sides of head
convex strongly. Occipital margin slightly concave. Eyes small and much convex. Antennal scape reaching occipital margin. Mesosoma convex strongly without suture. Anterodorsal portion of pronotum marginated. Propodeal spine triangular, slightly longer thanbroad at base. Petiole slightly longer than broad. Postpetiole slightly broader than length, and slightly broader than petiole. Headwith thick and diverging rugae. Mesosoma with thick rugae. Constricted anterior portion of pronotum with transverse rugae― 28 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

posteriorly. Petiole with longitudinal rugae. Body black, mandibles, antennal scapes, femora and tibiae brown, funiculus, and tarsi
reddish brown.
Remarks. We have not been able to examine type material of M. butteli.
Distribution. INDONESIA: Sumatra.

Myrmecina celebensis sp. nov.
(Fig. 10)
Holotype worker. TL 2.18, HL 0.53, HW 0.48, CI 91, SL 0.42, SI 87, PW 0.33, ML 0.56.
Paratype worker. TL 2.31, HL 0.54, HW 0.50, CI 93, SL 0.40, SI 81, PW 0.34, ML 0.59 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus not concave; median portion ofanterior margin feebly projected without a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions.
Anterior dorsal surface of labrum with paired small denticles, close to each other. Frontal carinae virtually absent, indistinguishable
from rugae on dorsum of head. Eyes small circularly and moderately convex, varying in size with maximum diameter 0.05-0.06 mm
and 6-9 ommatidia; malar space twice as long as diameter of eye or longer in profile; distance between occipital margin and posterior
margin of eye four times as long as diameter of eye or longer. Antennal scape short, not reaching posterolateral corner of head; antennal flange weakly developed.
Dorsal outline of mesosoma more or less flattened in profile. Pronotum without denticles; anterior portion marginate; anterior
ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection relatively broad. Eumetanotal spine
present. Propodeal spine triangular, longer than broad at base, extending over vertical posteriormost limit of propodeum in profile.
Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, apart from margin by its diameter. Petiole short,slightly longer than high in profile, longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process
variously developed from weakly raised median longitudinal ridge to distinct projection. Postpetiole broader than petiole, 1.5 times
as broad as its length except for helcium in dorsal view; anterolateral portion rounded; dorsal outline flattened in profile; ventral
outline projected rectangularly with two tops.
Anterior margin of gaster not concave.
Head with thin and weakly waved rugae; ventrolateral portion with longitudinal or oblique rugae weakly. Clypeus smooth andshining. Mesosoma with thin and longitudinal rugae. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae.
First gastral tergum punctured distinctly dorsally. First gastral sternum rugulose at anterior and lateral portion. Head with dense andshort pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole
and postpetiole without hairs on ventral surface. Head and mesosoma black to dark reddish brown, petiole and postpetiole reddish
brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Sulawesi, Tengah, nr. Morowali, Ranu River Area, 27. i-20. iv. 1980 (M. J. D. Brendell) (BMNH).
Paratype. 1 worker with same data as holotype (BMNH).
― 29 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 10. Myrmecina celebensis sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

Remarks. M. celebensis does not belong to any species complex. Among those species, M. celebensis can be distinguished from theremains by having the anterior clypeal margin without processes and the propodeal spine longer than broad at base.
Distribution. INDONESIA: Sulawesi.

― 30 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Myrmecina curvispina Zhou, Huang & Ma
(Fig. 11)Myrmecina curvispina Zhou, Huang & Ma, 2008: 286. Holotype worker, CHINA: Maoershan Natural Reserve, Xing’an County,
Guangxi Province (25°54’ N, 110°30’ E), August 22, 1998 (Shanyi Zhou). [Insect Collection, College of Life Sciences, Guangxi
Normal University, Guilin, China]. [not examined].

abcFig. 11. Myrmecina curvispina Zhou, Huang & Ma. a, dorsal view of mesosoma, petiole
and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view ofhead.

― 31 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Workers. TL 2.82-3.25, HL 0.67-0.77, HW 0.66-0.77, CI 98-102, SL 0.52-0.61, SI 77-83, PW 0.43-0.54, ML 0.70-0.83 (11measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin concave or straight; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth frequently unclear; apical tooth completely fused with preapical tooth; apical tooth about twice as long as preapical tooth. Dorsal surface of clypeus strongly concave;median portion of anterior margin projected and concave with distinct three processes; lateral portion simple, lacking sharp ridge infront of antennal insertions. Anterior dorsal surface of labrum with paired distinct denticles. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes small and not convex, varying in size with maximum diameter 0.06-0.11 mm and4-6 ommatidia. Antennal scape short, just reaching posterolateral corner of head; antennal flange developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticle on dorsolateral portion; anterior portion marginate; anterior ventrolateral portion with denticle directing forward and downward. Furrow between pronotum and mesoepisternal
projection more or less broad. Eumetanotal spine present but small. Propodeal spine elongate, directing backward, extending oververtical posteriormost limit of propodeum in profile; apex of spines curved upward distinctly. Propodeal lobe low. Propodeal spiracle
large, situated near posterior margin of propodeum, apart from margin by its diameter. Petiole short, slightly longer than high in
profile, and slightly longer than broad in dorsal view; small but distinct dorsal crest located at midlength in profile; subpetiolar processsmall, usually forming acute anterior apex. Postpetiole slightly broader than petiole in dorsal view; dorsal outline flattened or slightly
convex in profile; ventral outline distinctly projected with acute anterior apex.
Anterior margin of gaster concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with thin rugae longitudinally; ventrolateral portion smooth and shining. Dorsal surface of mandible sometimes appears
paired yellow spots. Clypeus smooth and shining. Mesosoma with thin rugae longitudinally on dorsal and lateral portion. Forecoxasmooth and shining. Petiole and postpetiole with a few distinct rugae. First gastral tergum punctured or rugulose weakly dorsally.
First gastral sternum smooth and shining. Head with sparse and long pilosity on dorsum. Mesosoma with sparse and long pilosity on
dorsum, hairs of pronotum slightly shorter than propodeal spine. Dorsal pilosity of petiole as long as that of mesosoma. Petiole
without hairs on ventral surface. Postpetiole with 2-4 hairs on ventral surface. Body black, mandibles, antennae and legs yellowish
brown, forecoxae reddish brown.
Other material examined. VIETNAM: Cuc Phuong N. P., Nho Quan Dist., Ninh Binh Prov., 11. viii. 1998, VN98-HO-022 (H.
Okido) (IEBR); Cuc Phuong N. P., Nho Quan Dist., Ninh Binh Prov., 11. viii. 1998, VN98-HO-021 (IEBR, KUEC, KUIC); Tam Dao,
900-1240 m alt., Vinh Phuc Prov. (Sk. Yamane).
Remarks. M. curvispina forms to the M. gracilis complex comprising M. cuvispina, M. gracilis sp. nov. and M. raviwonghei. The
definition and the distribution of the M. gracilis complex are given in the latter section. Among those species, M. curvispina can be
distinguished from the remains by having the smooth and shining first gastral sternum.
Distribution. VIETNAM: Vinh Phuc Prov., Ninh Binh Prov.

Myrmecina dasynota sp. nov.
(Fig. 12)― 32 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Holotype worker. TL 2.31, HL 0.56, HW 0.51, CI 91, SL 0.45, SI 88, PW 0.35, ML 0.59.
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin flattened; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 4 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin feebly projected without a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles, close to each other. Frontal carinae present but very short,abcFig. 12. Myrmecina dasynota sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

― 33 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

running back to the level of the posterior margin of the eyes. Eyes extremely small circularly and moderately convex with maximum
diameter 0.05 mm, with 6 ommatidia; malar space twice as long as diameter of eye or longer in profile; distance between occipitalmargin and posterior margin of eye four times as long as diameter of eye or longer. Antennal scape short, just reaching posterolateralcorner of head; antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion marginate;
anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present
but much reduced and unclear. Propodeal spine only angulately projected, not spine-like in profile. Propodeal lobe low. Propodeal
spiracle very large, situated near posterior margin of propodeum, distance between posterior margin of spiracle and posterior marginof propodeum shorter than diameter of spiracle. Petiole long, longer than high in profile, and longer than broad in dorsal view; dorsal
crest absent; subpetiolar process absent. Postpetiole slightly broader than petiole in dorsal view; dorsal outline largely convex inprofile; ventral outline slightly projected with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with irregular rugae which are thin; ventrolateral portion feebly sculptured but shining. Clypeus smooth and shining.
Mesosoma with waved rugae which are thin and longitudinal. Forecoxa smooth and shining. Petiole and postpetiole weakly sculptured. First gastral segment smooth and shining. Head with extremely dense, and relatively long pilosity on dorsum. Mesosoma withextremely dense, and relatively long pilosity on dorsum, hairs of pronotum much longer than propodeal spine. Petiole without hairson ventral surface. Postpetiole with 8 hairs on ventral surface. Body yellow.
Holotype worker, MALAYSIA: Sabah, Gn. Silam, 620 m, 1983 (R. Leakey) (BMNH).
Remarks. M. dasynota does not belong to any species complex. The species is a distinctive species in having the yellow body color,the extremely dense pilosity and the small eye comprising about 10 ommatidia.
Distribution. MALAYSIA: Borneo (Sabah).

Myrmecina dechai sp. nov.
(Fig. 13)
Holotype worker. TL 2.62, HL 0.59, HW 0.62, CI 105, SL 0.46, SI 74, PW 0.43, ML 0.67.
Paratype workers. TL 2.68-2.69, HL 0.61-0.62, HW 0.62, CI 100-103, SL 0.46-0.48, SI 74-77, PW 0.42, ML 0.59-0.67 (2measured).
Worker. Head subrectangular, as long as or slightly longer than broad in full-face view; median portion of occipital margin flattened;
occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth
tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave;median portion of anterior margin slightly projected with a small median process or three processes; lateral portion simple, lacking
sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, which are relatively
well separated. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes small and usually convex
moderately, varying in size with maximum diameter 0.07-0.08 mm and 12-13 ommatidia; malar space twice as long as diameter of
eye or shorter in profile; distance between occipital margin and posterior margin of eye four times as long as diameter of eye. Anten― 34 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 13. Myrmecina dechai sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

nal scape short, not reaching posterolateral corner of head; antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion marginate;
anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection usually broad sometimes narrow.
Eumetanotal spine present but small. Propodeal spine triangular, as long as broad at base, feebly extending over vertical posteriormost
limit of propodeum in profile; tips directing outward in dorsal view. Propodeal lobe low. Propodeal spiracle small, situated nearposterior margin of propodeum, distance between posterior margin of spiracle and posterior margin of propodeum variable, as long as
diameter of spiracle or longer. Petiole short, as long as high in profile, slightly longer than broad in dorsal view; dorsal crest absent;― 35 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

subpetiolar process absent. Postpetiole slightly broader than petiole in dorsal view; dorsal outline flattened in profile; ventral outline
slightly projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head weakly punctured with longitudinal rugae diverging posteriorly on occipital portion, and transverse rugae running alongoccipital carina; ventrolateral portion with longitudinal rugae weakly, but shining. Clypeus smooth and shining. Mesosoma weakly
punctured with transverse rugae on anterior portion, and longitudinal rugae on posterior portion. Forecoxa almost smooth andshining. Petiole and postpetiole with distinct longitudinal rugae. First gastral segment smooth and shining. Head with dense andshort pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum as long as propodeal spine. Petiole
without hairs on ventral surface. Postpetiole with 2 hairs on ventral surface. Body and forecoxae black to reddish brown, mandibles,
antennae and legs yellowish brown.
Holotype worker, THAILAND: Kaeng Krachan N. P., 17. xi. 85 (I. Löbl & D. Burckhardt) (BMNH).
Paratypes. 2 workers with same data as holotype; 2 workers with same data as holotype but 19. xi. 85 (BMNH).
Remarks. M. dechai does not belong to any species complex. But in terms of the mesosoma with transverse rugae and relative lengthof the antennal scape, M. dechai is similar to M. poringensis sp. nov. The similarity of those species does not always reflect the relationships, because the characters mentioned above tend to change independently. Among those species, M. dechai can be distinguished from the remains by having the triangular propodeal spine pointing backward.
Distribution. THAILAND: Phetchaburi Prov.

Myrmecina dolichothrix sp. nov.
(Fig. 14)
Holotype worker. TL 2.74, HL 0.62, HW 0.61, CI 97, SL 0.58, SI 95, PW 0.44, ML 0.72.
Paratype workers. TL 2.60-2.75, HL 0.58-0.63, HW 0.58-0.61, CI 96-101, SL 0.54-0.61, SI 93-100, PW 0.42-0.45, ML 0.69-0.72 (5measured).
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin slightly concave;
occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth
tooth); apical tooth strong, third tooth robust, followed by 4 small teeth and a small basal tooth; small teeth frequently unclear. Dorsal
surface of clypeus not concave; median portion of anterior margin projected and flattened without a median process; lateral portion
simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles very small but
distinct. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and convex, varying in sizewith maximum diameter 0.10-0.11 mm and 6-7 ommatidia; malar space much twice as long as diameter of eye or shorter in profile;
distance between occipital margin and posterior margin of eye three times as long as diameter of eye or shorter. Antennal scape long,extending beyond posterolateral corner of head; antennal flange developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior portion marginate; anterior ventrolateral
portion angulate. Mesoepisternal projection not developed; furrow between pronotum and mesoepisternal projection relatively broad.
Eumetanotal spine absent. Propodeal spine elongate, 1.5 times as long as broad at base, extending over vertical posteriormost limit― 36 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 14. Myrmecina dolichothrix sp. nov. a, dorsal view of mesosoma, petiole
and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, fullface view of head.

of propodeum in profile. Propodeal lobe usually low, but sometimes raised. Propodeal spiracle large, situated near posterior marginof propodeum, distance between posterior margin of spiracle and posterior margin of propodeum shorter than diameter of spiracle.
Petiole long, slightly longer than high in profile, and longer than broad in dorsal view; very small dorsal crest located at midlength inprofile; subpetiolar process absent, but median longitudinal ridge at ventral portion present. Postpetiole slightly broader than petiole
in dorsal view; lateral margin straight; dorsal outline flattened in profile; ventral outline projected rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
― 37 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIHead with straight rugae; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma with oblique rugae
on dorsum. Forecoxa smooth and shining. Petiole and postpetiole with a few distinct rugae. First gastral segment smooth andshining. Head with sparse and long pilosity on dorsum. Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum
slightly longer than propodeal spine. Dorsal pilosity of petiole as long as that of mesosoma. Petiole without hairs on ventral surface.
Postpetiole usually without hairs on ventral surface. Body black or dark reddish brown, mandibles, antennae and legs yellowish
brown to reddish brown.
Holotype worker, PHILIPPINES: Surigao del N, SW Bacuag, Payapag, Little Baguio Waterfall, 6. ii. 2000 (S. Schödl) (NHMW).
Paratypes. 5 workers with same data as holotype (NHMW).
Remarks. M. dolichothrix does not belong to any species complex. But in terms of the body size, head shape, body sclupture, andthe shape of the propodeal spine, M. dolichothrix is similar to M. gopa sp. nov. The similarity of those species does not always reflect
the relationships, because the characters mentioned above tend to change independently. Among those species, M. dolichothrix canbe distinguished from the remains by having the long pilosity and the anterior clypeal margin without processes.
Distribution. PHILIPPINES: Mindanao.

Myrmecina elegans sp. nov.
(Fig. 15)
Holotype worker. TL 3.42, HL 0.84, HW 0.72, CI 86, SL 0.69, SI 96, PW 0.48, ML 0.94.
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth. Dorsal surface of clypeus concave; median portion of anterior margin slightly
projected with a median process or three processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior
dorsal surface of labrum with paired small denticles, which are relatively well separated. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes extremely small circularly and moderately convex with maximum diameter 0.06 mm,with 8 ommatidia; malar space distinctly twice as long as diameter of eye or longer in profile; distance between occipital margin andposterior margin of eye distinctly four times as long as diameter of eye or longer. Antennal scape short, just reaching posterolateralcorner of head; antennal flange fully developed.
Mesosoma slender in profile. Pronotum without denticles on anterior dorsolateral portion; anterior portion not marginate; anterior ventrolateral portion not angulate. Mesoepisternal projection much reduced; furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present. Propodeal spine elongate, much longer than broad at base, extending over verticalposteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine,distance between posterior margin of spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole long,
longer than high in profile, and much longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process
present, bearing acute anterior apex. Postpetiole expanded posteriorly as broad as petiole in dorsal view; dorsal outline flattened in
profile; ventral outline slightly projected rectangularly without acute anterior apex.
Anterior margin of gaster slightly concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
― 38 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 15. Myrmecina elegans sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Head with irregular rugae which are thin; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma
feebly sculptured, but almost smooth and shining. Forecoxa smooth and shining. Petiole and postpetiole smooth and shining. First
gastral segment smooth and shining. Head with dense and long pilosity on dorsum. Mesosoma with dense and long pilosity on
dorsum, hairs of pronotum slightly shorter than propodeal spine. Petiole with 2 hairs on ventral surface. Postpetiole with 8 hairs on
ventral surface. Body yellowish brown.

― 39 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Holotype worker, MALAYSIA: Sabah, Kinabalu, 1880 m, 26. iv. 87 (Burckhardt & Löbl) (BMNH).
Remarks. M. elegans does not belong to any species complex. But M. elegans is similar to M. aspera sp. nov. on the basis of somecharacters (see the remarks of M. aspera). Among those species, M. elegans can be distinguished from the remains by having theelongate propodeal spine, the smooth and shining dorsum of the pronotum, and the eye comprising 10 ommatidia or less.
Distribution. MALAYSIA: Borneo (Sabah).

Myrmecina glabra sp. nov.
(Fig. 16)
Holotype worker. TL 2.06, HL 0.50, HW 0.46, CI 94, SL 0.38, SI 83, PW 0.34, ML 0.53.
Paratype worker. TL 2.04, HL 0.50, HW 0.48, CI 95, SL 0.38, SI 80, PW 0.34, ML 0.51 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 4 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin slightly projected and concave; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior
dorsal surface of labrum with paired denticles small and blunt, close to each other. Frontal carinae present but very short andunclearly, running back to the level of the posterior margin of the eyes. Eyes very small and moderately convex, varying in size withmaximum diameter 0.06-0.08 mm and 11-13 ommatidia; malar space two times as long as diameter of eye in profile; distance between
occipital margin and posterior margin of eye four times as long as diameter of eye. Antennal scape short, not reaching posterolateralcorner of head; antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion not marginate; anterior ventrolateral portion slightly projected. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal
spine present but small. Propodeal spine triangular, as long as broad at base, feebly extending over vertical posteriormost limit ofpropodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, apart from marginby its diameter. Petiole short, slightly longer than high in profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process much reduced; ventral outline slightly concave. Postpetiole broader than petiole in dorsal view;lateral margin almost straight; dorsal outline flattened in profile; ventral outline projected with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head completely smooth and shining on dorsum and ventrolateral portion. Clypeus smooth and shining. Mesosoma completely
smooth and shining. Forecoxa smooth and shining. Petiole and postpetiole with weak irregular rugae. First gastral segment smoothand shining. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum
as long as or slightly shorter than propodeal spine. Petiole and postpetiole without hairs on ventral surface. Body black to reddish
brown, mandibles, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Neg. Sembilan, Pasoh For. Res., xi. 1994 (M. Brendell, K. Jackson & S. Lewis) (BMNH).
Paratype. 1 worker with same data as holotype (BMNH).

― 40 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 16. Myrmecina glabra sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Remarks. M. glabra does not belong to any species complex. This species is easily distinguished from the other species by the completely smooth, shining mesosoma in dorsal view.
The species inhabits in litter.
Distribution. MALAYSIA: The Malay Pen.

― 41 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Myrmecina gopa sp. nov.
(Fig. 17)
Holotype worker. TL 2.61, HL 0.61, HW 0.58, CI 95, SL 0.53, SI 92, PW 0.41, ML 0.67.
Paratype workers. TL 2.54-2.90, HL 0.57-0.65, HW 0.54-0.62, CI 91-97, SL 0.50-0.56, SI 88-96, PW 0.37-0.44, ML 0.64-0.75 (25measured).
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin slightly concave;
occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth
tooth); apical tooth strong, third tooth robust, followed by 4-5 small teeth and a small basal tooth; small teeth frequently unclear.
Dorsal surface of clypeus slightly concave; median portion of anterior margin projected with three processes; lateral portion simple,
lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles very small. Frontal carinae
virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and moderately convex, varying in size with maximum
diameter 0.08-0.11 mm and 5-6 ommatidia; malar space twice as long as diameter of eye or shorter in profile; distance between occipital margin and posterior margin of eye three times as long as diameter of eye. Antennal scape long, extending beyond posterolateralcorner of head; antennal flange developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior portion usually marginate; anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection more or less broad. Eumetanotal spine absent.
Propodeal spine elongate, 1.5 times as long as broad at base, extending over vertical posteriormost limit of propodeum in profile.
Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, distance between posterior margin of
spiracle and posterior margin of propodeum shorter than diameter of spiracle. Petiole short, as long as or slightly longer than high in
profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process present at anterior
portion, usually bearing acute anterior apex, but varied in shape, sometimes triangular or lobed with a hole. Postpetiole broader thanpetiole in dorsal view; lateral margin usually straight; anterior portion sharply raised; ventral outline projected with two acute points.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae distinctly; ventrolateral portion usually smooth and shining. Clypeus smooth and shining. Mesosoma
with straight rugae distinctly. Forecoxa smooth and shining. Petiole and postpetiole with a few rugae. First gastral segment smoothand shining. Head with relatively dense and long pilosity on dorsum. Mesosoma with dense and long pilosity on dorsum, hairs ofpronotum slightly shorter than propodeal spine. Dorsal pilosity of petiole slightly longer than that of mesosoma. Petiole without hairson ventral surface. Postpetiole usually without hairs on ventral surface. Head and mesosoma black, petiole, postpetiole and gasterreddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Kebun Raya, Bogor, W. Java, 11. ix. 1996, FI96-375 (F. Ito) (BZM).
Paratypes. numerous workers, ergatogynes and males with same data as holotype; numerous workers, ergatogynes and males withsame data as holotype but 17-20. ii. 1993, FI93-157; numerous workers and ergatogynes, and 1 male with same data as holotype but
17-20. ii. 1993, FI93-159; numerous workers, ergatogynes and males with same data as holotype but 17-20. ii. 1993, FI93-162;
numerous workers, ergatogynes and males with same data as holotype but 17-20. ii. 1993, FI93-163 (BZM, KUEC, KUIC).
Remarks. M. gopa does not belong to any species complex. But M. gopa is similar to M. dolichothrix sp. nov. on the basis of somecharacters (see the remarks of M. dolichothrix). Among those species, M. gopa can be distinguished from the remains by having theshort pilosity and the anterior clypeal margin with three processes. This species corresponds to “Myrmecina sp. A” of Ito (1996),― 42 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 17. Myrmecina gopa sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Aoki & Ito (1997), Ito et al. (2001) and Tsuji et al. (2001).
Colony of this species is unicolonial, because their nest are contiguously rather than uniformly distributed, and conspecific nonnestmates are easily incorporated into alien nests (Tsuji et al., 2001). Ito (1996) reported that this species, which has a specialized
myrmecophilous oribatid mite Aribates javensis Aoki, Takaku & Ito (Ito & Takaku, 1994; Aoki et al., 1994), had no queen, but ergatogyne as a reproductive caste. Ergatogyne distinguish from worker only in width and length of first gastral tergum. Colonies of thisspecies are polygynous, having 8 ergatogynes on average per colony, and all mated ergatogynes reproduce without aggressiveinteractions.
― 43 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIDistribution. INDONESIA: Java.

Myrmecina gracilis sp. nov.
(Fig. 18)
Holotype worker. TL 2.53 (excluding mandible length), HL 0.66, HW 0.56, CI 85, SL 0.50, SI 89, PW 0.43, ML 0.66.
Paratype workers. TL 2.30-2.90, HL 0.55-0.70, HW 0.50-0.62, CI 86-95, SL 0.38-0.51, SI 75-88, PW 0.37-0.48, ML 0.59-0.75 (9measured).
Worker. Head subrectangular, longer than broad in full-face view; lateral margin more or less straight; median portion of occipitalmargin distinctly concave; occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength(third small tooth or sixth tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth
and a basal tooth unclear. Dorsal surface of clypeus distinctly concave; median portion of anterior margin slightly projected andconcave without median process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface oflabrum with paired small denticles distinctly. Frontal carinae present, running to near occipital corners; frequently indistinguishable
from rugae on dorsum of head. Eyes very small and moderately convex, varying in size with maximum diameter 0.05-0.06 mm and4-7 ommatidia. Antennal scape short, not reaching posterolateral corner of head; antennal flange not developed.
Dorsal outline of mesosoma flattened or slightly convex in profile. Pronotum with very small denticles in dorsolateral portion;
anterior portion not marginate; anterior ventrolateral portion usually not angulate, but sometimes angulate. Eumetanotal spine present
but small. Propodeal spine elongate and weakly curved upward, extending over vertical posteriormost limit of propodeum in profile.
Propodeal lobe low. Propodeal spiracle usually large, situated near posterior margin of propodeum, distance between posterior marginof spiracle and posterior margin of propodeum shorter than diameter of spiracle. Petiole slightly long, slightly longer than high in
profile, and longer than broad in dorsal view; midpoint of both lateral margins concave in dorsal view, but varying; small but distinct
dorsal crest located at midlength in profile; anterior portion of subpetiolar process much developed with acute anterior apex. Postpetiole broader than petiole in dorsal view; anterior portion sometimes sharply raised in profile; ventral outline slightly projected.
Anterior margin of gaster concave in dorsal view; first gastral sternum forming a median longitudinal ridge distinctly on anterior
central surface.
Head punctured with thin rugae longitudinally; ventrolateral portion punctured with thin rugae. Clypeus smooth and shining.
Mesosoma punctured with thin rugae parallely. Forecoxa smooth and shining. Petiole and postpetiole punctured with unclear irregular rugae. First gastral tergum punctured distinctly. First gastral sternum feebly rugulose or punctured. Head with dense and shortpilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Longest
dorsal pilosity of petiole as long as propodeal spine. Petiole without hairs on ventral surface. Postpetiole without hairs on ventral
surface. Body black or dark reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Sayap Kinabalu, ca. 1000 m alt., Sabah, Borneo (K. Eguchi) (UMS).
Paratypes. 4 workers, MALAYSIA: Sarawak, 4th Division, Gn. Mulu NP., 8. ii. 78 or ii. 78 or v-vii. 1978 (N. M. Collins; P. M.
Hammond & J. E. Marshall); 1 worker, Sabah, Kinabalu, 1500 m, 25. iv. 87 (Burckhardt & Löbl); 2 workers, Sabah, Crocker Ra. NP.,
KK-Tambunan, 1270 m, 17. v. 87 (Burckhardt & Löbl); 1 worker, Sabah, Kibongol Valley, 7 km N. Tambunan (Löbl & Burckhardt);
2 workers, Sabah, Gn. Silam, 620 m, 1983 (R. Leakey) (BMNH).

― 44 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 18. Myrmecina gracilis sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Other material examined. INDONESIA: Teluk Kabah, Kutai N. P., E. Kalimantan (Sk. Yamane) (KUIC).
Remarks. M. gracilis belongs to the M. gracilis complex comprising M. curvispina, M. gracilis and M. raviwonghei. The definition
and the distribution of the gracilis complex are given in the latter section. Among those species, M. gracilis can be distinguished fromthe remains by having the sculptured first gastral sternum with ridge at anterior median portion, and the long head. The species inhabits in soil.
Distribution. MALAYSIA: Borneo (Sabah, Sarawak); INDONESIA: Borneo (Kalimantan).
― 45 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Myrmecina grandis sp. nov.
(Fig. 19)
Holotype worker. TL 3.47, HL 0.86, HW 0.75, CI 87, SL 0.70, SI 94, PW 0.50, ML 0.91.
Paratype workers. TL 3.30-3.60, HL 0.79-0.90, HW 0.74-0.82, CI 88-93, SL 0.70-0.75, SI 91-98, PW 0.47-0.51, ML 0.90-0.96 (11measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible straight; apical tooth strong, third tooth robust, followed by 6
small teeth. Dorsal surface of clypeus not concave; median portion of anterior margin much projected with distinct three processes;
lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles,
close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes medium-sized and moderately convex, varying in size with maximum diameter 0.11-0.13 mm and 7-8 ommatidia; malar space twice as long as diameter of
eye in profile; distance between occipital margin and posterior margin of eye three times as long as diameter of eye or longer. Antennal scape short, just reaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma convex in profile. Pronotum with denticles on dorsolateral portion, but variable, sometimes reduced;
anterior portion marginate; anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection
narrow. Eumetanotal spine present. Propodeal spine triangular, slightly longer than broad at base, extending over vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance
between posterior margin of spiracle and posterior margin of propodeum much longer than diameter of spiracle. Petiole short, as longas or slightly longer than high in profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar
process usually present, bearing acute anterior apex. Postpetiole broader than petiole and expanded anteriorly in dorsal view; anterior
portion sharply raised in profile; ventral outline slightly projected rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head usually with straight rugae distinctly, sometimes weakly waved; ventrolateral portion smooth and shining. Clypeus smoothand shining. Mesosoma with straight rugae longitudinally. Forecoxa smooth and shining. Petiole and postpetiole with a few rugae.
First gastral segment smooth and shining. Head with relatively dense and long pilosity on dorsum. Mesosoma with dense and longpilosity on dorsum, hairs of pronotum slightly shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with
4-8 hairs on ventral surface. Head and mesosoma black, antennal scape, petiole, postpetiole and gaster reddish brown, mandibles,
antennal funiculus and legs yellowish brown to brown.
Holotype worker, INDONESIA: Ulu Gadut, Padang, W-Sumatra, 17. viii. 1996, 19. ii. 1997 fixed, FI96-379 (K. Ohkawara) (BZM).
Paratypes. numerous workers and ergatogynes with same data as holotype (BZM, KUEC, KUIC, UMS).
Other material examined. INDONESIA: Ulu Gadut nr Padang, W. Sumatra (Sk. Yamane) (KUIC). MALAYSIA: Sepilok Forest,
Sabah, Borneo, (K. Eguchi) (KUEC); Poring, Sabah, Borneo, (T. Kikuta) (KUEC); Poring Hot Springs, Sabah (Burckhardt & Löbl)(BMNH); 4th Division Gn. Mulu NP., Sarawak (P. M. Hammond & J. E. Marshall) (BMNH); Ulu Gombak, ca. 250 m alt., Serangor
Prov., Malay Pen. (Sk. Yamane) (KUIC). Brunei: Tasek Merimbum, (K. Eguchi) (KUEC).
Remarks. M. grandis does not belong to any species complex. The species can be distinguished from other species by having thestraight masticatory margin of the mandible, simple anterior margin of the smooth and shining first gastral tergum, the short antennal― 46 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

a

b

d

c
Fig. 19. Myrmecina grandis sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, masticatory
margin of mandible.

scape just reaching occipital corner, the much projected anterior clypeal margin with distinct three processes, and the medium-sized
eye. This species corresponds to “Myrmecina sp. E” of Aoki & Ito (1997). Aoki & Ito (1997) reported that this species had a myrmecophilous oribatid mite Protoribates myrmecophilus Aoki & Ito, which was also found in the nest of M. maryatiae sp. nov. (=Myrmecina sp. B). Ergatogynes are present in this species.
Distribution. MALAYSIA: Borneo (Sabah, Sarawak), The Malay Pen.; BRUNEI: Tasek Merimbum; INDONESIA: Sumatra.
― 47 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Myrmecina guangxiensis Zhou
(Fig. 20)
Myrmecina guangxiensis Zhou, 2001: 139, 237. Holotype worker, CHINA: Huashuichong Natural Reserve, Hezhou City, Guangxi,
1.ix.1998 (Shanyi Zhou). Paratype worker, Da Ping Shan Natural Reserve, 18.ix.1998 (John Fellowes). [not seen].
Worker. TL 2.22, HL 0.56, HW 0.54, CI 97, SL 0.48, SI 88, PW 0.38, ML 0.59.
ergatogyne. TL 2.72, HL 0.61, HW 0.61, CI 100, SL 0.53, SI 87, PW 0.45, ML 0.67.
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin almost flattened; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 4-5 small teeth and a stout basal tooth bearing blunt apex. Dorsal surface of clypeus
concave; median portion of anterior margin weakly projected without a median process; lateral portion raised into a sharp ridge ofshield wall on each side, in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles, fused with each otherat base. Frontal carinae present, running back to or beyond the level of the posterior margin of the eyes. Eyes very small, varying insize with maximum diameter 0.06 mm and 4-5 ommatidia. Antennal scape long, extending beyond posterolateral corner of head;antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior ventrolateral portion not angulate. Eumetanotal spine present but small. Propodeal spine triangular, directing upward and backward, not reaching vertical posteriormost limit ofpropodeum in profile; tips directing outward in dorsal view. Propodeal lobe low. Propodeal spiracle large, situated near base of
propodeal spine, apart from margin by its diameter. Petiole short; small dorsal crest located slightly at anterior portion in profile;subpetiolar process projecting forward with acute apex. Postpetiole slightly broader than petiole in dorsal view; lateral margin straightand not marginated; dorsal outline flattened in profile; ventral outline projecting forward with small but distinct apex.
Anterior margin of gaster concave in dorsal view.
Head with thin and irregular rugae and punctures weakly, but variable; ventrolateral portion smooth and shining, without rugae.
Mandibles with large yellow spots on dorsal surface. Clypeus weakly punctured or smooth and shining. Mesosoma with thick rugae
irregularly in profile. Mesosoma with thick rugae wavy and divergent anteriorly (forming “Y”-shape). Petiole and postpetiole witha few longitudinal rugae. First gastral tergum smooth and shining dorsally. Head with sparse pilosity on dorsum. Mesosoma withsparce pilosity on dorsum. Pilosity of pronotum more or less longer than propodeal spines. Pilosity of petiole and postpetiole relatively longer than that of mesosoma. Postpetiole with hairs on ventral surface. Head, mesosoma, petiole and postpetiole varying fromreddish brown to black, legs yellowish, forecoxae yellowish brown, gaster reddish brown.
Other material examined. VIETNAM: Cuc Phuong, 320 m alt., Ninh Binh Prov., 2. vi. 1998 (S. Nomura) (KUEC); Cuc Phuong,
340 m alt., Ninh Binh Prov., 1. vi. 1998 (KUEC); Dong Dong Bai, 170 m alt., Cuc Phuong, Ninh Binh Prov. (S. Nomura).
Remarks. M. guangxiensis forms the M. vieti complex. The other members of the complex are: M. nomurai sp. nov. and M. vieti sp.
nov. Among those species, M. guangxiensis can be distinguished from the remains by having the short and sculptured petiole which
is not expanded laterally at the anterior portion in dorsal view, and the mesosoma with the rugae diverging anteriorly. Some morphological variations, such as the smooth and shining clypeus and the more strongly punctured mesosoma, are found in this species.
Ergatogynes are present in this species.
The species inhabits in soil.
― 48 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 20. Myrmecina guangxiensis Zhou. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Distribution. VIETNAM: Ninh Binh Prov.

Myrmecina gymnocephala sp. nov.
(Fig. 21)
Holotype worker. TL 2.78, HL 0.67, HW 0.66, CI 99, SL 0.62, SI 93, PW 0.43, ML 0.74.
Paratype workers. TL 2.71-2.93, HL 0.66-0.67, HW 0.66-0.68, CI 100-104, SL 0.61-0.63, SI 90-93, PW 0.42-0.45, ML 0.70-0.80 (5― 49 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 21. Myrmecina gymnocephala sp. nov. a, dorsal view of mesosoma,
petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin flattened or slightly concave;
occipital corners widely rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or
sixth tooth); apical tooth strong, third tooth robust, followed by 4-5 small teeth and a stout basal tooth bearing blunt apex; small teeth
frequently unclear. Dorsal surface of clypeus slightly concave; median portion of anterior margin weakly projected with a medianprocess but the process frequently unclear; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal
surface of labrum with paired small denticles distinctly, closed to each other. Frontal carinae present but variable in length; running― 50 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

back to or beyond the level of the posterior margin of the eyes. Eyes extremely large and convex, varying in size with maximum
diameter 0.14-0.16 mm and 7-8 ommatidia; malar space as long as diameter of eye in profile. Antennal scape long, extending beyond
posterolateral corner of head distinctly; antennal flange weakly developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior ventrolateral portion not angulate.
Eumetanotal spine absent. Propodeal spine triangular and very small, not reaching vertical posteriormost limit of propodeum in
profile. Propodeal lobe slightly raised. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior
margin of spiracle and posterior margin of propodeum slightly longer than diameter of spiracle. Petiole long, longer than high in
profile, and longer than broad in dorsal view; small but distinct dorsal crest located almost at midlength in profile; subpetiolar processabsent or unclear; ventral outline flattened or slightly convex. Postpetiole broader than petiole in dorsal view; anterolateral marginrounded; dorsal outline slightly convex or flattened in profile; ventral outline projected rectangularly at anterior and posterior portion.
Anterior margin of gaster not concave in dorsal view.
Head smooth and shining, completely absent any sculpture except for frontal carinae and a few rugae in front of eyes; ventrolateral portion smooth and shining. Mandibles smooth and shining, frequently with yellow spots on dorsal surface. Clypeus smooth andshining. Mesosoma with with longitudinal rugae which are thick, sometimes median portion smooth and shining in dorsal view.
Petiole and postpetiole with weak rugae. Head with sparse pilosity on dorsum. Mesosoma with sparse pilosity on dorsum. Pilosityof pronotum slightly shorter than maximum diameter of eye. Pilosity of petiole and postpetiole as long as that of mesosoma. Petiole
without hairs on ventral surface. Postpetiole with about 4 hairs on ventral surface. Head, mesosoma, petiole and postpetiole reddish
brown.
Holotype worker, INDONESIA: N. C. Seram, Manusela N. P., Wae Mual Plain, 25. vii-9. xi. 1987 (M. J. D. Brendell) (BMNH).
Paratypes. 7 workers with same data as holotype (BMNH).
Other material examined. PHILIPPINES: 28 km S Calapan, Balete, Mindoro, (Schillhammer); 10 km W Puerto Galera, Mindoro(Schillhammer) (NHMW).
Remarks. M. gymnocephala belongs to the M. macrops complex comprising M. breviata sp. nov., M. gymnocephala and M. macrops
sp. nov. This species is different from the related species in having the developed propodeal spine, and the pronotum and mesonotumwith straight rugae. Dorsal surface of the mesosoma of the workers distributed in Seram is sculptured only on the lateral portion,
while that of the workers distributed in Mindoro is sculptured on the entire surface.
Distribution. PHILIPPINES: Mindoro; INDONESIA: Seram.

Myrmecina inflata sp. nov.
(Fig. 22)
Holotype worker. TL 3.18, HL 0.79, HW 0.75, CI 95, SL 0.67, SI 89, PW 0.42, ML 0.80.
Paratype workers. TL 3.12-3.26, HL 0.74-0.84, HW 0.74-0.80, CI 92-101, SL 0.66-0.72, SI 83-98, PW 0.46-0.51, ML 0.78-0.86 (7measured).
Worker. Occipital margin of head strongly concave in full-face view; lateral margin strongly convex; dorsal outline strongly convex― 51 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

in profile; occipital corners projected posteriorly. Masticatory margin of mandible straight; apical tooth strong, third tooth weakly
developed, followed by 6 small teeth; small teeth frequently unclear. Dorsal surface of clypeus not concave; median portion of anterior margin weakly projected; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface oflabrum with paired small denticles distinctly, which are well separated. Frontal carinae present, running back to occipital corners, butvery obscure, difficult to distinguish from rugae on dorsum of head. Eyes large and convex, varying in size with maximum diameter
0.14-0.18 mm and 6-8 ommatidia. Antennal scape short, just reaching posterolateral corner of head; antennal flange developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles on dorsolateral portion; anterior ventrolateralabcFig. 22. Myrmecina inflata sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

― 52 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

portion forming small denticle or an angle. Eumetanotal spine present but small and unclear. Propodeal spine triangular, directing
upward and backward, just reaching vertical posteriormost limit of propodeum in profile; tips directing outward in dorsal view. Propodeal lobe low. Propodeal spiracle medium-sized, situated near base of propodeal spine, distance between posterior margin of
spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole shorter than high in profile; dorsal crest locatedat midlength in profile; ventral outline flattened; subpetiolar process present. Postpetiole as broad as petiole and expanded posteriorly
in dorsal view; dorsal outline slightly convex or flattened in profile; ventral outline projected with anterior apex at anterior portion.
Anterior margin of gaster concave in dorsal view.
Head with thin rugae which are weakly waved; ventrolateral portion with 2-3 rugae curved downward. Clypeus smooth andshining. Mesosoma with irregular rugae in profile. Mesosoma with thick rugae distinctly. Petiole and postpetiole with distinct rugae.
Head and mesosoma with dense pilosity on dorsum, hairs of pronotum longer than propodeal spine. Pilosity of petiole and postpetiole
slightly longer than that of mesosoma. Postpetiole with about 4 hairs, but varies on ventral surface. Body black, mandibles, antennae
and legs reddish brown to yellowish brown.
Holotype worker, THAILAND: Phang Nga Prov., Tone Chong-Fah Waterfall, 20 km S. Takuapa, 100-200 m H, 11-14. 01. 1996 (A.
Schulz, K. Vock) (NHMW).
Paratypes. 8 workers with same data as holotype (NHMW).
Remarks. M. inflata forms to the M. spinosa complex comprising M. inflata, M. mahuana sp. nov., M. spinosa sp. nov., M. magnificens and M. tridentata sp. nov. The definition and the distribution of the spinosa complex are given in the latter section. Among thosespecies, M. inflata can be distinguished from the remains by having the concave anterior margin of the first gastral tergum and thestraight masticatory margin of the mandible.
Distribution. THAILAND: Phang Nga Prov.

Myrmecina insulana sp. nov.
(Fig. 23)
Holotype worker. TL 2.33, HL 0.56, HW 0.51, CI 91, SL 0.43, SI 84, PW 0.38, ML 0.61.
Paratype workers. TL 2.22-2.37, HL 0.53-0.58, HW 0.51-0.53, CI 92-97, SL 0.43-0.45, SI 82-88, PW 0.37-0.40, ML 0.58-0.59 (4measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus not concave; median portion ofanterior margin feebly projected with or without median paired processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, relatively close to each other. Frontal carinae
virtually absent, indistinguishable from rugae on dorsum of head. Eyes very small circularly and usually convex moderately, varying
in size with maximum diameter 0.06-0.08 mm and 10-11 ommatidia; malar space twice as long as diameter of eye or longer in profile;
distance between occipital margin and posterior margin of eye distinctly four times as long as diameter of eye or longer. Antennal
scape short, not or just reaching posterolateral corner of head; antennal flange fully developed.
― 53 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Dorsal outline of mesosoma convex in profile. Pronotum with or without denticle; anterior portion marginate; anterior ventrolateral portion projected. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present but small.
Propodeal spine triangular, variable in length, usually extending over vertical posteriormost limit of propodeum in profile. Propodeal
lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior margin of spiracle and posterior
margin of propodeum longer than diameter of the spiracle. Petiole short, as long as or slightly longer than high in profile, slightly
longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process variously developed from weakly

a

b

d

c
Fig .23. Myrmecina insulana sp. nov. a, dorsal view of mesosoma, petiole and postpetiole;b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, ventrolateral portion of head.

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

raised median longitudinal ridge to distinct projection. Postpetiole slightly broader than petiole in dorsal view; dorsal outline flattenedor slightly convex in profile; ventral outline projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view.
Head with wavy rugae which are thin; ventrolateral portion with transverse, or vertical rugae sometimes unclearly. Clypeus
smooth and shining. Mesosoma with wavy rugae which are thick. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastral segment smooth and shining. Head with dense and short pilosity on dorsum. Mesosoma with dense andshort pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with
2-4 hairs on ventral surface. Head and mesosoma black, petiole and postpetiole reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Mahua Waterfall area, ca. 1000 m alt., Crocker Range N. P., Sabah, 5. xi. 2000 (J. Fellowes) (UMS).
Paratypes. 2 workers and 1 queen with same data as holotype; 2 workers with same data as holotype but (H. Okido); MALAYSIA:
Sabah, Kinabalu N. P., Poring Hot Springs, East Ridge, 560 m, 17. viii. 95 (C. Brühl); Danum Valley, Sabah, Borneo, 29. viii. 1995
(H. Okido); Sayap Kinabalu, ca. 1000 m alt., Sabah, Borneo, 14. vii. 1996 (Sk. Yamane); Sayap Kinabalu, ca. 1000 m alt., Sabah,
Borneo, 14. vii. 1996, S-17 (K. Eguchi) (UMS, KUEC, KUIC).
Other material examined. MALAYSIA: Crocker Ra., Sabah (Burckhardt & Löbl) (BMNH); Poring Hot Springs, Sabah (Burckhardt& Löbl) (BMNH); 4th Division, Gn. Mulu NP. (N. M.Collins; B. Bolton) (BMNH); Marudi, Sarawak (P. Hammond) (BMNH); Neg.
Sembilan, Pasoh For. Res. (M. Brendell, K. Jackson & L. Ficken) (BMNH). INDONESIA: Teluk Kabah, Kutai N. P., E. Kalimantan(Sk. Yamane); Mt. Gede, Cibodas, W. Java (F. Ito); Sukarami nr. Padang, W. Sumatra (F. Ito).
Remarks. M. insulana does not belong to any species complex. But in terms of the eye size, shape and sculpture of the first gastralsegment, relative length of the antennal scape, and sculpture of the mesosoma, M. insulana is similar to M. sabahna sp. nov. and M.
yamanei sp. nov. The similarity of those species does not always reflect the relationships, because the characters mentioned above
tend to change independently. Among those species, M. insulana can be distinguished from the remains by having ventrolateral
portion of the head with transverse rugae. The species inhabits in soil and litter.
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); INDONESIA: Sumatra, Borneo (Kalimantan), Java.

Myrmecina inthanonensis sp. nov.
(Fig. 24)
Holotype worker. TL 3.38, HL 0.80, HW 0.78, CI 97, SL 0.70, SI 91, PW 0.51, ML 0.88.
Paratype worker. TL 3.39, HL 0.80, HW 0.74, CI 92, SL 0.70, SI 96, PW 0.50, ML 0.90 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin distinctly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus concave; median portionof anterior margin projected with a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior
dorsal surface of labrum with paired small denticles distinctly, close to each other. Frontal carinae virtually absent, indistinguishable― 55 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 24. Myrmecina inthanonensis sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

from rugae on dorsum of head. Eyes slightly large and distinctly convex, varying in size with maximum diameter 0.10-0.14 mm and7-8 ommatidia; malar space twice as long as diameter of eye or slightly longer in profile; distance between occipital margin and posterior margin of eye four times as long as diameter of eye or longer. Antennal scape long, slightly extending beyond posterolateralcorner of head; antennal flange fully developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion feebly marginate; anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine
present but small, somtimes reduced. Propodeal spine triangular, slightly longer than broad at base, not or just reaching vertical― 56 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, apart from margin by its diameter. Petiole short, as long as or slightly longer than high in profile, and as long as or slightly
longer than broad in dorsal view; dorsal crest located anteriorly in profile; subpetiolar process absent, but median longitudinal ridge
at ventral portion present. Postpetiole expanded posteriorly and slightly broader than petiole and larger than petiole in dorsal view;dorsal outline flattened in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head punctured with thick rugae distinctly; ventrolateral portion punctured with thick rugae distinctly. Clypeus smooth andshining. Mesosoma punctured with thick rugae distinctly. Forecoxa with transverse rugae weakly, but sometimes smooth andshining. Petiole and postpetiole with thick rugae. First gastral segment smooth and shining. Head with relatively sparse and shortpilosity on dorsum. Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum as long as propodeal spine. Petiole with
4 hairs on ventral surface. Postpetiole with 4-6 hairs on ventral surface. Body and forecoxae black, mandibles, antennae and legs
yellowish brown to reddish brown.
Holotype worker, THAILAND: Doi Inthanon, 7. xi. 85 (Löbl & Burckhardt) (BMNH). Paratypes. 5 workers with same data as holotype (BMNH).
Remarks. M. inthanonensis does not belong to any species complex. But in terms of the petiole with the subpetiolar process ormedian longitudinal ridge, relative length of the antennal scape, the mesosoma with the eumetanotal spine and relative size of the
postpetiole, M. inthanonensis is similar to M. nigra sp. nov. The similarity of those species does not always reflect the relationships,
because the characters mentioned above tend to change independently. Among those species, M. inthanonensis can be distinguishedfrom the remains by having the distinctly large postpetiole comparing with the petiole, the anterior clypeal margin with a single
median process, and the mesosoma with straight and parallel rugae on dorsal surface.
Distribution. THAILAND: Chiang Mai Prov.

Myrmecina itoi sp. nov.
(Fig. 25)
Holotype worker. TL 3.41, HL 0.82, HW 0.76, CI 93, SL 0.64, SI 84, PW 0.56, ML 0.93.
Paratype workers. TL 3.30-3.46, HL 0.78-0.82, HW 0.70-0.77, CI 90-94, SL 0.62-0.67, SI 88-89, PW 0.50-0.54, ML 0.93-0.96 (2measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin slightly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible straight; apical tooth strong, third tooth robust, followedby 6 small teeth. Dorsal surface of clypeus concave; median portion of anterior margin distinctly projected without a median process;lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired blunt denticles, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and convex,
varying in size with maximum diameter 0.16-0.18 mm and 9-10 ommatidia; malar space 1.5 times as long as diameter of eye or longer
in profile; distance between occipital margin and posterior margin of eye twice as long as diameter of eye or longer. Antennal scapeshort, just reaching posterolateral corner of head; antennal flange weakly developed.
― 57 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 25. Myrmecina itoi sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateralview of mesosoma, petiole and postpetiole; c, full-face view of head.

Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion marginate, but unclearly;
anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection more or less broad. Eumetanotal
spine present but small. Propodeal spine triangular, as long as broad at base, just reaching vertical posteriormost limit of propodeum
in profile. Propodeal lobe low. Propodeal spiracle slightly large, situated near base of propodeal spine, distance between posterior
margin of spiracle and posterior margin of propodeum much longer than diameter of spiracle. Petiole short, as long as high in profile,and slightly longer than broad in dorsal view; small dorsal crest located at midlength in profile; subpetiolar process present at anterior
portion, bearing acute anterior apex. Postpetiole slightly broader than petiole and expanded posteriorly in dorsal view; dorsal outline― 58 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

slightly convex or flattened in profile; ventral outline projected with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with irregular rugae distinctly; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma withwavy rugae distinctly. Forecoxa smooth and shining. Petiole and postpetiole with a few rugae. First gastral segment smooth andshining. Head with relatively dense and short pilosity on dorsum. Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum as long as or slightly shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 2-4 hairs on
ventral surface. Body black, mandibles, antennal funiculus and legs yellowish brown to reddish brown, antennal scapes dark reddish
brown.
Holotype worker, MALAYSIA: Ulu Gombak, 24. x. 1996, FI96-695 (F. Ito) (UMS).
Paratypes. 11 workers and 2 males with same data as holotype (UMS, KUEC, KUIC).
Other material examined. MALAYSIA: Tawau Hills N. P., Sabah, Borneo, (Sk. Yamane) (KUIC). INDONESIA: Ketambe, Leuser
N. P., Aceh, N-Sumatra (D. Iskandar) (KUEC).
Remarks. M. itoi does not belong to any species complex. The species can be distinguished from other species by having the straightmasticatory margin of the mandible, simple anterior margin of the smooth and shining first gastral tergum, the short antennal scapejust reaching occipital corner, the long petiole, and the large eye.
This species, which inhabits in rotten wood, is found without specialized myrmecophilous oribatid mite (Ito, per. comm.).
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra.

Myrmecina lambirensis sp. nov.
(Fig. 26)
Holotype worker. TL 2.84 (head separated), HL 0.68, HW 0.61, CI 89, SL 0.56, SI 92, PW 0.38, ML 0.82.
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin slightly concave;
occipital corners rounded, not projected posteriorly. Masticatory margin of mandible straight; apical tooth strong, third tooth robust,
followed by 6 small teeth. Dorsal surface of clypeus slightly concave; median portion of anterior margin much projected with distinct
three processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum withpaired small denticles distinctly. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes mediumsized and moderately convex with maximum diameter 0.10 mm, with 6 ommatidiain the longest row; malar space twice as long as
diameter of eye in profile; distance between occipital margin and posterior margin of eye three times as long as diameter of eye or
longer. Antennal scape short, just reaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion not marginate; anterior
ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection relatively narrow. Eumetanotal spine
present but small. Propodeal spine triangular, length as long as broad at base, not reaching vertical posteriormost limit of propodeum
in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior margin of
spiracle and posterior margin of propodeum much longer than diameter of spiracle. Petiole short, length as long as high in profile,― 59 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 26. Myrmecina lambirensis sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

longer than broad in dorsal view; dorsal crest located slightly at anterior portion in profile; subpetiolar process reduced. Postpetiole
slightly broader than petiole; lateral margin straight; anterior portion slightly raised in profile; ventral outline slightly projected
rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with weakly waved rugae; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma with weakly
waved rugae. Forecoxa smooth and shining. Petiole and postpetiole with a few rugae. First gastral segment smooth and shining.
Head with relatively sparse and short pilosity on dorsum. Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum― 60 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

longer than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 4 hairs on ventral surface. Head and mesosoma black, antennal scapes, petiole, postpetiole and gaster reddish brown, mandibles, antennal funiculus and legs yellowish brown
to reddish brown.
Holotype worker, MALAYSIA: Bt. Lambir, Lambir N. P., Miri, Sarawak, 22. viii. 1995 (H. Okido) (UMS).
Remarks. M. lambirensis does not belong to any species complex. The species can be distinguished from other species by having thesmooth and shining dorsal median portion of the mesosoma and lateral portion of the pronotum, the anterior clypeal margin with
distinct three processes, and the short antennal scape.
Distribution. MALAYSIA: Borneo (Sarawak).

Myrmecina lombokensis sp. nov.
(Fig. 27)
Holotype worker. TL 2.45, HL 0.58, HW 0.58, CI 100, SL 0.48, SI 83, PW 0.42, ML 0.64.
Paratype worker. TL 2.65, HL 0.61, HW 0.61, CI 100, SL 0.53, SI 87, PW 0.43, ML 0.62 (1 measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth unclear. Dorsal surface of clypeus concave;median portion of anterior margin projected with a median process that are very small; lateral portion simple, lacking sharp ridge infront of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly. Frontal carinae virtually absent,
indistinguishable from rugae on dorsum of head. Eyes large and convex with maximum diameter 0.08 mm, with 5 ommatidia; malar
space twice as long as diameter of eye in profile; distance between occipital margin and posterior margin of eye three times as long as
diameter of eye. Antennal scape short, just reaching posterolateral corner of head; antennal flange weakly developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion marginate; anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present but small; length
shorter than broad at base. Propodeal spine elongate, directing backward and weakly curved upward, extending over vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum,apart from the margin by its diameter. Petiole short, as long as high in profile, and longer than broad in dorsal view; small but distinct
dorsal crest located at midlength in profile; subpetiolar process small with acute anterior apex clearly. Postpetiole broader than petiole
and expanded anteriorly in dorsal view; dorsal outline flattened or slightly convex in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head punctured with thin rugae longitudinally; ventrolateral portion shining, but punctured. Clypeus smooth and shining.
Mesosoma punctured with thin rugae longitudinally. Forecoxa smooth and shining. Petiole and postpetiole with a few distinct rugae.
First gastral segment smooth and shining. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on
dorsum, hairs of pronotum shorter than apical third segment of antenna. Petiole without hairs on ventral surface. Postpetiole with 2
hairs on ventral surface. Body black to dark reddish brown, mandibles, antennae and legs brown to yellowish brown.

― 61 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 27. Myrmecina lombokensis sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Holotype worker, INDONESIA: Pusuk Pass, 260 m alt., Lombok I., S 08°28´xE 116°5´, 1. x. 2000, TUS KO-3 (K.Ogata) (KUEC).
Paratype. 1 worker, INDONESIA: Senaru, 750 m alt., Lombok I., S 08°20´xE 118°24´, 30. ix. 2000, TUS KO-2 (K. Ogata) (KUEC).
Remarks. M. lombokensis does not belong to any species complex. But M. lombokensis is similar to M. asthena sp. nov. on the basisof some characters (see the remarks of M. asthena). Among those species, M. lombokensis can be distinguished from the remains by
having the medium-sized and ovally eye comprising about 14 ommatidia.

― 62 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Distribution. INDONESIA: Lombok.

Myrmecina longiseta sp. nov.
(Fig. 28)
Holotype worker. TL 3.37, HL 0.80, HW 0.76, CI 95, SL 0.72, SI 95, PW 0.56, ML 0.93.
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin slightly projected with a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions.
Anterior dorsal surface of labrum with paired small denticles distinctly, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes slightly large and distinctly convex with maximum diameter 0.11 mm, with 7 ommatidia; malar space twice as long as diameter of eye in profile; distance between occipital margin and posterior margin of eye four times
as long as diameter of eye or longer. Antennal scape long, slightly extending beyond posterolateral corner of head; antennal flange
fully developed.
Dorsal outline of mesosoma convex in profile. Pronotum with unclear denticles on dorsolateral portion; anterior portion marginate; anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine
present but small. Propodeal spine triangular, longer than broad at base, not reaching vertical posteriormost limit of propodeum in
profile. Propodeal lobe low. Propodeal spiracle small, situated near base of propodeal spine, apart from margin by its diameter.
Petiole long, longer than high in profile, and slightly longer than broad in dorsal view; dorsal crest located at midlength in profile;subpetiolar process and median longitudinal ridge at ventral portion absent. Postpetiole as broad as petiole in dorsal view; dorsal
outline flattened or slightly convex in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae which are thin distinctly; ventrolateral portion with longitudinal rugae anteriorly, and smooth andshining posteriorly. Clypeus smooth and shining. Mesosoma with some transverse rugae anteriorly, and longitudinal rugae posteriorly. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastral segment smooth and shining. Headwith sparse and long pilosity on dorsum. Mesosoma with sparse and extremely long pilosity on dorsum, hairs of pronotum twice aslong as propodeal spine or longer. Petiole without hairs on ventral surface. Postpetiole with 2 hairs on ventral surface. Head, mesosoma, forecoxae, petiole and postpetiole black, gaster dark reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Sumatra, Padang (no collector’s name) (BMNH).
Remarks. M. longiseta does not belong to any species complex. But M. longiseta is similar to M. boltoni sp. nov., M. monticola sp.
nov. and M. parallela sp. nov. on the basis of some characters (see the remarks of M. boltoni). Among those species, M. longiseta canbe distinguished from the remains by having the sparse and extremely long pronotal pilosity and the short propodeal spine.
Distribution. INDONESIA: Sumatra.

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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 28. Myrmecina longiseta sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Myrmecina macrops sp. nov.
(Fig. 29)
Holotype worker. TL 2.80, HL 0.66, HW 0.69, CI 105, SL 0.56, SI 81, PW 0.48, ML 0.75.
Paratype workers. TL 2.57-2.77, HL 0.62-0.67, HW 0.65-0.70, CI 104-105, SL 0.53-0.59, SI 81-86, PW 0.45-0.50, ML 0.69-0.73 (5measured).

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin flattened or slightly concave;
occipital corners widely rounded, not projected posteriorly in profile. Masticatory margin of mandible bent at midlength (third small
tooth or sixth tooth); apical tooth strong, third tooth robust, followed by about 4 small teeth and a stout basal tooth bearing blunt apex;
small teeth unclear. Dorsal surface of clypeus slightly concave; median portion of anterior margin weakly projected with a medianprocess; the process frequently unclear; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal
surface of labrum with paired small denticles distinctly, very closed to each other without fusion. Frontal carinae present, runningback to about mid point level of eyes but very unclearly. Eyes extremely large and moderately convex, varying in size with maximum
diameter 0.18 mm and 8-9 ommatidia; malar space as long as diameter of eye in profile. Antennal scape long, extending beyondabcFig. 29. Myrmecina macrops sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

posterolateral corner of head; antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior ventrolateral portion angulate. Eumetanotal
spine present but small, frequently unclear. Propodeal spine triangular, just reaching vertical posteriormost limit of propodeum in
profile. Propodeal lobe low. Propodeal spiracle small, situated near base of propodeal spine, distance between posterior margin of
spiracle and posterior margin of propodeum much longer than diameter of spiracle. Petiole short, as long as high in profile, andslightly longer than broad in dorsal view; small but distinct dorsal crest located almost at midlength in profile; subpetiolar process
projecting forward with acute apex. Postpetiole slightly broader than petiole in dorsal view; lateral margin straight; dorsal outline
slightly convex or flattened in profile; ventral outline projected rectangularly, frequently undeveloped.
Anterior margin of gaster not concave in dorsal view.
Head distinct foveolate, but variable from opaque to foveolate on the whole surface; ventrolateral portion smooth and shining.
Mandibles smooth and shining sometimes with yellow spots on dorsal surface. Clypeus smooth and shining. Mesosoma with wavy
rugae distictly. Petiole and postpetiole with wavy longitudinal rugae. Head with more or less dense pilosity on dorsum. Mesosoma
with more or less dense pilosity on dorsum. Pilosity of pronotum slightly shorter than propodeal spine, but variable in length, sometimes longer than propodeal spine. Pilosity of petiole and postpetiole longer than that of mesosoma. Petiole with or without hairs on
ventral surface. Postpetiole with about 4 hairs on ventral surface. Body dark reddish brown.
Holotype worker, MALAYSIA: Sepilok Forest, Sabah, Borneo, 4. vii. 1998 (K. Eguchi) (UMS). Paratypes. 9 workers and 1 queen(dealate) with same data as holotype (KUIC, KUEC).
Other material examined. MALAYSIA: Danum Valley, Sabah (C. Brühl); Deramakot Forest Reserve, Sabah (C. Brühl); Poring, 600
m alt., Kinabalu Park, Sabah (T. Kikuta); Tambunan Village Resort Centre, Sabah (H. Okido); Ulu Gombak (F. Ito; Sk. Yamane).
INDONESIA: Brastagi, 1500 m, N. Sumatra (Rougemont); Padang, Sumatra (no data); Kebun Raya, Bogor, W-Java (F. Ito).
Remarks. M. macrops belongs to the M. macrops complex comprising M. breviata sp. nov., M. gymnocephala sp. nov. and M.
macrops. This species is different from the related species in having developed propodeal spine, the pronotum and mesonotum withwavy rugae. This species corresponds to “Myrmecina sp. M” of Ito et al. (2001).
The species inhabits in soil and litter, and has been collected by the pitfall trap.
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra, Java.

Myrmecina magnificens Wong & Guénard
(Fig. 30)
Myrmecina magnificens Wong & Guénard, 2016: 132. Holotype worker, SINGAPORE: Seletar Trail, Central Catchment Nature
Reserve, 1.395141°N, 103.802595°E, 47 m, 2.ix.2015, leaf litter (MKL Wong), label “MW020915-L1.1” (ANTWEB1009004)(LKCNHM). Paratypes. Four workers (ANTWEB1009005–ANTWEB1009008), same data as holotype (SBSHKU). [notexamined].
Workers. TL 4.02-4.26, HL 1.08-1.13, HW 1.06-1.09, CI 96-99, SL 0.93-1.01, SI 87-93, PW 0.69, ML 1.10-1.14 (5 measured).

― 66 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 30. Myrmecina magnificens Wong & Guénard. a, dorsal view of mesosoma, petiole
and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view ofhead.

Worker. Occipital margin strongly concave in full-face view; occipital corners projected posteriorly. Masticatory margin of mandible
straight; apical tooth strong, third tooth acute, followed by 6 small but distinct teeth. Dorsal surface of clypeus concave; median
portion of anterior margin weakly projected, without a median process; lateral portion without a sharp ridge of shield wall, but weakly
carinate on each side, in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles, close to each otherwithout fusion. Frontal carinae present, running to near occipital corners but frequently unclear. Eyes large and convex, varying insize with maximum diameter 0.18-0.20 mm and 8-10 ommatidia; malar space longer than diameter of eye in profile. Antennal scapejust reaching posterolateral corner of head; antennal flange developed.
― 67 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Dorsal outline of mesosoma slightly convex in profile. Pronotum with two or four small denticles on anterior dorsolateral portionin dorsal view, but frequently unclear; anterior ventrolateral portion with small denticle directing forward and downward but varying
in shape and direction; frequently not forming denticle but angulately. Eumetanotal spine long with acute apex. Propodeal spine
elongate, variable in length, strongly curved upward. Propodeal lobe low. Propodeal spiracle situated near base of propodeal spine,distance between posterior margin of spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole short,
almost as long as high in profile; small dorsal crest located anterioriy in profile; subpetiolar process triangular but variable; ventral
outline flattened. Postpetiole almost as broad as petiole and expanded posteriorly in dorsal view; lateral portion not marginated; dorsal
outline slightly convex in profile; ventral outline projected at anterior portion.
Anterior margin of gaster concave in dorsal view.
Head with weakly waved rugae which are thick and longitudinal; ventrolateral portion usually with longitudinal rugae which are
curved at posterior portion. Mandibles sometimes with reddish brown spots on dorsal surface clearly. Clypeus smooth and shining.
Mesosoma with weakly waved rugae which are thick and longitudinal; lateral portion with weakly waved rugae which are thick.
Petiole and postpetiole with a few longitudinal rugae. First gastral tergum smooth and shining. Head and mesosoma with long pilosity on dorsum. Pilosity of pronotum as long as propodeal spines or longer. Head, mesosoma, petiole and postpetiole black, legs
varying from yellowish brown to light reddish brown, forecoxae varying from reddish brown to black, gaster reddish brown.
Other material examined. MALAYSIA: Tioman Is., 17-22. ix. 1992, MT-1 (F. Ito) (UMS); Tioman Is., 17-22. ix. 1992, MT-2 (F.
Ito); Tioman Is., 17-22. ix. 1992, MT-3 (F. Ito) (KUIC, KUEC); Ulu Gombak (F. Ito); Pasoh (K. Masuko). INDONESIA: Sitiung,
W-Sumatra (F. Ito).
Remarks. M. magnificens belongs to the M. spinosa complex. The other members of the complex are: M. inflata sp. nov., M.
mahuana sp. nov., M. spinosa sp. nov. and M. tridentata sp. nov. The definition and the distribution of the spinosa complex are givenin the latter section. Among those species, M. magnificens can be distinguished from the remains by having the concave anterior
margin of the first gastral tergum, the bent masticatory margin of the mandible, and the strongly elevated propodeal spine.
Ergatogynes are present in the species, and only distinguished from the workers in width and length of the first gastral tergum(Ito, 1996).
The species inhabits in rotten branches (Ito, pers. comm.), and litter.
Distribution. MALAYSIA: Tioman Is., The Malay Pen.; INDONESIA: Sumatra.

Myrmecina mahuana sp. nov.
(Fig. 31)
Holotype worker. TL 3.73, HL 0.96, HW 0.94, CI 98, SL 0.93, SI 98, PW 0.61, ML 1.02.
Worker. Head convex on lateral margin; occipital margin strongly concave in full-face view; occipital corners projected posteriorly.
Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical tooth strong, third tooth weakly developed,
followed by 5 small teeth and a stout basal tooth bearing blunt apex; small teeth frequently unclear. Dorsal surface of clypeus notconcave; median portion of anterior margin weakly projected with a median process bearing two tops; lateral portion simple, lacking
sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles, close to each other. Frontal― 68 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 31. Myrmecina mahuana sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

carinae present, running back to or beyond the level of the posterior margin of the eyes, but very obscure, difficult to distinguish fromrugae on dorsum of head. Eyes large and convex with maximum diameter 0.18 mm, with 11 ommatidia; malar space twice as long as
diameter of eye or shorter in profile. Antennal scape long, extending beyond posterolateral corner of head; antennal flange
developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior ventrolateral portion
with small denticle. Eumetanotal spine present but small and unclear. Propodeal spine triangular, directing upward and backward,
just reaching vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle small, situated near base― 69 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

of propodeal spine, between posterior margin of spiracle and posterior margin of propodeum much longer than diameter of spiracle.
Petiole longer than high in profile; dorsal crest absent; subpetiolar process absent. Postpetiole slightly broader than petiole in dorsal
view; lateral margin straight; dorsal outline slightly convex or flattened in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view.
Head with wavy rugae distinctly; ventrolateral portion with 2-3 rugae longitudinally. Mandibles smooth and shining. Clypeus
smooth and shining. Mesosoma with irregular rugae. Petiole and postpetiole with longitudinal rugae. Head and mesosoma with longpilosity on dorsum, hairs of pronotum longer than apical second segment of antenna. Pilosity of petiole and postpetiole as long as thatof mesosoma. Postpetiole with paired hairs on ventral surface. Head, mesosoma, petiole and postpetiole dark reddish brown, mandible yellowish brown, gaster black.
Holotype worker, MALAYSIA: Mahua Waterfall area, ca. 1000 m alt., Crocker Range N. P., Sabah (H. Okido) (UMS).
Other material examined. MALAYSIA: Gun. Lawit, Summit ridge, Trengganu (T. Clay) (BMNH). INDONESIA: Sumatra, Padang(no collector’s name) (BMNH).
Remarks. M. mahuana belongs to the M. spinosa complex. Among those species, M. mahuana can be distinguished from the remainsby having the propodeal spine directed posteriorly and simple anterior margin of the first gastral tergum.
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Sumatra.

Myrmecina maryatiae sp. nov.
(Fig. 32)
Holotype worker. TL 2.58, HL 0.62, HW 0.58, CI 92, SL 0.50, SI 86, PW 0.43, ML 0.66.
Paratype workers. TL 2.37-2.80, HL 0.58-0.67, HW 0.53-0.62, CI 90-97, SL 0.48-0.53, SI 84-94, PW 0.38-0.45, ML 0.59-0.74 (24measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus concave; median portion ofanterior margin slightly projected and concave without a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes extremely large and moderately convex, varying in size with maximum diameter
0.11-0.16 mm and 7-10 ommatidia; malar space as long as diameter of eye in profile; distance between occipital margin and posterior
margin of eye twice as long as diameter of eye. Antennal scape short, just reaching posterolateral corner of head; antennal flange fully
developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum usually without denticle, but sometimes with small denticles;
anterior portion marginate; anterior ventrolateral portion angulate; anterior dorsolateral portion angulate. Furrow between pronotum
and mesoepisternal projection broad. Eumetanotal spine present but small. Propodeal spine triangular, as long as broad at base, notor just reaching vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle small, situated near base― 70 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum much longer than diameter ofspiracle. Petiole short, as long as high in profile, and slightly longer than broad in dorsal view; dorsal crest located at midlength inprofile; subpetiolar process usually present with acute anterior apex. Postpetiole slightly broader than petiole in dorsal view; dorsal
outline flattened in profile; ventral outline slightly projected rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with irregular rugae which are thin distinctly; ventrolateral portion smooth and shining. Clypeus smooth and shining, but

a

b

d

c
Fig. 32. Myrmecina maryatiae sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, lateral portionof clypeus.

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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

posterior portion sculptured. Mesosoma with wavy rugae distinctly. Forecoxa smooth and shining. Petiole and postpetiole with a
few rugae. First gastral segment smooth and shining. Head with relatively dense and short pilosity on dorsum. Mesosoma with dense
and short pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole
with 2-4 hairs on ventral surface. Body usually black, mandibles, antennae and legs yellowish brown to black.
Holotype worker, MALAYSIA: Ulu Gombak, 27. x. 1996, FI96-721, 19. ii. 1997 fixed (F. Ito) (UMS).
Paratypes. numerous workers, queens and males with same data as holotype; 15 workers and 1 queen with same data as holotype
but vii-x. 1992, FI92MG502, no date for fix; numerous workers, queens and males with same data as holotype but vii-x. 1992,
FI92MG565, no date for fix; numerous workers, queens and males with same data as holotype but 24. x. 1996, FI96-694; 32 workers,
1 queen and 1 male with same data as holotype but 24. x. 1996, FI96-698; numerous workers, queens and males with same data as
holotype but 26. x. 1996, FI96-700; 4 workers, 1 queen and 2 males with same data as holotype but 28. x. 1996, FI96-729; numerous
workers, queen and male with same data as holotype but 28. x. 1996, FI96-738; numerous workers, queens and males with same data
as holotype but 28. x. 1996, FI96-740; 10 workers and 1 queen with same data as holotype but 18. vii. 1998, FI98-108, no date for fix(KUEC, KUIC, UMS).
Other material examined. MALAYSIA: Bunga Buah, nr Genting H. L. (Sk. Yamane) (KUIC, KUEC); Neg. Sembilan, Pasoh For.
Res. (M. Brendell, K. Jackson & L Ficken) (BMNH); 4th Division, Gn. Mulu N. P. (P. M. Hammond & J. E. Marshall) (BMNH);
Selangor, Gombak (B. Bolton) (BMNH); Tower Region, Lambir N. P., Miri, Sarawak (Sk. Yamane) (KUEC, KUIC); Niah N. P.,
Sarawak (Sk. Yamane) (KUEC); Danum Valley, Sabah, Borneo (H. Okido) (KUEC). BRUNEI: Tasek Merimbum, (K. Eguchi) (KUEC,
KUIC). INDONESIA: Sangkimah, Kutai N. P., E. Kalimantan (Sk. Yamane) (KUEC). THAILAND: Tone Chong-Fah Waterfall, 20
km S. Takuapa, Phang Nga Prov. (A. Schulz) (NHMW).
Remarks. M. maryatiae does not belong to any species complex. But M. maryatiae is similar to the species of the macrops complex
on the basis of two characters. Among those species, M. maryatiae can be distinguished from the remains by having the sculptured
head and the short antennal scape. This species corresponds to “Myrmecina sp. B” of Aoki & Ito (1997). Aoki & Ito (1997) reported
that this species had a myrmecophilous oribatid mite Protoribates myrmecophilus Aoki & Ito, which was also found in the nest of
M.grandis sp. nov. (=Myrmecina sp. E).
The species inhabits in litter or under bark.
Distribution. THAILAND: Phang Nga Prov.; MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); INDONESIA: Borneo(Kalimantan).

Myrmecina monticola sp. nov.
(Fig. 33)
Holotype worker. TL 3.13, HL 0.78, HW 0.72, CI 93, SL 0.69, SI 96, PW 0.51, ML 0.86.
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portion― 72 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

of anterior margin projected with four processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior
dorsal surface of labrum with small but distinct paired denticles, very close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes medium-sized and convex with maximum diameter 0.11 mm, with 8 ommatidia; malar
space twice as long as diameter of eye or shorter in profile; distance between occipital margin and posterior margin of eye three times
as long as diameter of eye or longer. Antennal scape long, extending beyond posterolateral corner of head; antennal flange weakly
developed.

abcFig. 33. Myrmecina monticola sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face viewof head.

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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion marginate;
anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present
but small. Propodeal spine triangular, as long as broad at base, not reaching vertical posteriormost limit of propodeum in profile.
Propodeal lobe low. Propodeal spiracle medium-sized, situated near posterior margin of propodeum, apart from margin by its diameter. Petiole short, slightly longer than high in profile, and slightly longer than broad in dorsal view; dorsal crest located at midlength
in profile; subpetiolar process and median longitudinal ridge at ventral portion absent. Postpetiole slightly broader than petiole andexpanded posteriorly in dorsal view; larger than petiole in profile; dorsal outline flattened; ventral outline slightly projected
rectangularly.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head punctured with thin rugae weakly; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma withthick rugae diverging anteriorly on dorsal surface. Anterior portion of pronotum with a few transverse rugae in dorsal view. Forecoxasmooth and shining. Petiole and postpetiole with thick rugae. First gastral segment smooth and shining. Head with relatively dense
and long pilosity on dorsum. Mesosoma with dense and long pilosity on dorsum, hairs of pronotum longer than propodeal spine.
Petiole without hairs on ventral surface. Postpetiole with 4 hairs on ventral surface. Body black, mandibles, antennae and legs yellowish brown to brown, forecoxae dark reddish brown.
Holotype worker, MALAYSIA: Cameron Highlands, 1500 m alt., Malay Pen., 25. iv. 2000 (K. Eguchi) (UMS).
Remarks. M. monticola does not belong to any species complex. But M. monticola is similar to M. boltoni sp. nov. on the basis ofsome characters (see the remarks of M. boltoni). Among those species, M monticola can be distinguished from the remains by having
the anterior clypeal margin with median paired processes and the smooth and shining forecoxa.
Distribution. MALAYSIA: The Malay Pen.

Myrmecina muluensis sp. nov.
(Fig. 34)
Holotype worker. TL 3.56, HL 0.85, HW 0.80, CI 94, SL 0.72, SI 90, PW 0.50, ML 0.99.
Paratype worker. TL 3.43, HL 0.84, HW 0.75, CI 90, SL 0.72, SI 96, PW 0.50, ML 0.93 (1 measured).
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin much concave; occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth);apical tooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave;median portion of anterior margin slightly projected with a median process or three processes; lateral portion simple, lacking sharp
ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles, which are well separated. Frontal carinae
virtually absent, indistinguishable from rugae on dorsum of head. Eyes small circularly and moderately convex, varying in size withmaximum diameter 0.09 mm and 12-13 ommatidia; malar space distinctly twice as long as diameter of eye or longer in profile; distance between occipital margin and posterior margin of eye five times as long as diameter of eye or longer. Antennal scape long,extending beyond posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion not― 74 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 34. Myrmecina muluensis sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face viewof head.

marginate; anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal
spine distinctly present. Propodeal spine elongate, much longer than broad at base, not reaching or just reaching vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance
between posterior margin of spiracle and posterior margin of propodeum longer than diameter of the spiracle. Petiole long, longer
than high in profile, and much longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process variously developed from weakly raised median longitudinal ridge to distinct projection. Postpetiole slightly broader than petiole in― 75 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

dorsal view; anterolateral margin rounded; anterior portion raised in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view.
Head with thin rugae longitudinally; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma withstraight rugae which are thick. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastral segmentsmooth and shining. Head with dense and long pilosity on dorsum. Mesosoma with dense and long pilosity on dorsum, hairs ofpronotum as long as propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 2-4 hairs on ventral surface. Head,mesosoma, petiole, postpetiole reddish brown or yellowish brown but variable, mandibles, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Sarawak, 4th Division, Gn. Mulu NP., 20. iii. 78 (N. M. Collins) (BMNH).
Paratypes. 2 workers with same data as holotype (BMNH).
Remarks. M. muluensis does not belong to any species complex. But in terms of the smooth and shining ventrolateral portion of thehead, the petiole with subpetiolar process or median longitudinal ridge, the mesosoma with the eumetanotal spine and relative lengthof the antennal scape, M. muluensis is similar to M. sulcata Emery. The similarity of those species does not always reflect the relationships, because the characters mentioned above tend to change independently. Among those species, M muluensis can be distinguishedfrom the remains by having the small eye comprising 15 ommatidia or less, and the long head. The species inhabits in soil.
Distribution. MALAYSIA: Borneo (Sarawak).

Myrmecina nesaea Wheeler
Myrmecina nesaea Wheeler, 1924: 247. Syntype queen, INDONESIA: Sebesi I. (Dammerman) (depository unknown) [not seen].
Remarks. Worker of this species is unknown.
Distribution. INDONESIA: Sebesi I. (near Krakatau Is.)

Myrmecina nigra sp. nov.
(Fig. 35)
Holotype worker. TL 3.20, HL 0.77, HW 0.74, CI 96, SL 0.70, SI 96, PW 0.50, ML 0.82.
Paratype workers. TL 3.05-3.36, HL 0.76-0.80, HW 0.74-0.75, CI 94-98, SL 0.67-0.69, SI 91-92, PW 0.49-0.51, ML 0.77-0.79 (4measured).
Worker. Head subrectangular, as long as broad in full-face view or longer; median portion of occipital margin concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus concave; median portion ofanterior margin projected with three processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior
dorsal surface of labrum with paired small denticles, close to each other. Frontal carinae virtually absent, indistinguishable from rugae― 76 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 35. Myrmecina nigra sp. nov. a, dorsal view of mesosoma, petiole and postpetiole;b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

on dorsum of head. Eyes small circularly and moderately convex, varying in size with maximum diameter 0.08-0.10 mm and 13-14
ommatidia; malar space twice as long as diameter of eye in profile; distance between occipital margin and posterior margin of eye fourtimes as long as diameter of eye. Antennal scape long, extending beyond posterolateral corner of head; antennal flange weakly
developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior portion marginate but unclearly in dorsal
view; anterior ventrolateral portion more or less angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present but small, sometimes unclear. Propodeal spine triangular, extending over vertical posteriormost limit of― 77 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior
margin of spiracle and posterior margin of propodeum slightly longer than diameter of spiracle. Petiole short, as long as or slightly
longer than high in profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process variously developed from weakly raised median longitudinal ridge to distinct projection. Postpetiole slightly longer than petiole andexpanded posteriorly in dorsal view; anterior portion not raised in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view.
Head with wavy rugae which are thin distinctly; ventrolateral portion sculptured irregularly. Clypeus smooth and shining.
Mesosoma with wavy rugae distinctly. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastralsegment smooth and shining. Head with relatively dense and long pilosity on dorsum. Mesosoma feebly dense and long pilosity on
dorsum, hairs of pronotum as long as propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 4 hairs on ventral
surface. Body black to dark reddish brown, antennae and legs brown to yellowish brown.
Holotype worker, VIETNAM: Deo O Quy Ho, 1750 m alt., Lao Cai Prov., 28. vi. 1997 (S. Nomura) (KUEC).
Paratypes. 4 workers with same data as holotype (KUEC, IEBR, KUIC).
Remarks. M. nigra does not belong to any species complex. But M. nigra is similar to M. inthanonensis sp. nov. on the basis of somecharacters (see the remarks of M. inthanonensis). Among those species, M nigra can be distinguished from the remains by having theslightly large postpetiole comparing with the petiole, the anterior clypeal margin with three processes, and the mesosoma with wavy
and diverging rugae on dorsal surface.
Distribution. VIETNAM: Lao Cai Prov.

Myrmecina nitidiuscula Satria & Yamane
Myrmecina nitidiuscula Satria & Yamane, 2019: 188. Holotype worker, INDONESIA: Sumatra, Aceh, Leuser Ecosystem, Putri
Betung, 21.ix.2012 (R. Satria) (SEMUT20180326B) (MZB). Paratypes. Two workers (SEMUT20180808A, SEMUT20180808B),
same data as holotype (RSC); 2 workers (SKYUSI-FOR004-005), same locality, but at. >1100 m, good forest, 20.ix.2012 (Yamane &Syaukani) (SKYC); 1 dealate queen (SEMUT20181112), same locality, but alt. 850 m, plantation, 19.ix.2012 (Yamane & Syaukani)(RSC). [one SKYC paratype worker examined].
Remarks. M. nitidiuscula belongs to the macrops complex comprising M. breviata sp. nov., M. gymnocephala sp. nov. and M.
macrops sp. nov. This species is different from the related species in having the smooth and shingy head, the developed propodeal
spine, and the pronotum and mesonotum with straight rugae.
Distribution. INDONESIA: Sumatra.

Myrmecina nomurai sp. nov.
(Fig. 36)

― 78 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Holotype worker. TL 2.24, HL 0.56, HW 0.53, CI 94, SL 0.43, SI 82, PW 0.35, ML 0.60.
Paratype worker. TL 2.34, HL 0.58, HW 0.53, CI 90, SL 0.46, SI 88, PW 0.35, ML 0.63 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin moderately concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth acute, followed by 5 small teeth and a stout basal tooth bearing blunt apex. Dorsal surface of clypeus slightly
concave; median portion of anterior margin weakly projected with small median process but frequently unclear; lateral portion raised

a

b

d

c
Fig. 36. Myrmecina nomurai sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, antennal
flange.

― 79 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

into a sharp ridge of shield wall on each side, in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles
fused with each other at base. Frontal carinae present and running back to or beyond the level of the posterior margin of the eyes, but
frequently unclear. Eyes very small, varying in size with maximum diameter 0.06 mm and 6-7 ommatidia. Antennal scape short, justreaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma more or less flattened in profile. Pronotum without denticles; anterior ventrolateral portion notangulate. Eumetanotal spine present but small. Propodeal spine triangular, directing slightly upward and backward, not reachingvertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal
spine, apart from margin by its diameter. Petiole short; small dorsal crest located almost at midlength in profile; subpetiolar process
projecting forward with acute apex. Postpetiole much broader than petiole and usually expanded anteriorly in dorsal view; lateral
portion strongly marginated; dorsal outline flattened in profile; ventral outline projected forward with acute apex at anterior portion.
Anterior margin of gaster concave in dorsal view.
Head weakly punctured with irregular rugae which are thin, but variable; ventrolateral portion weakly punctured or smooth andshining, without rugae. Mandibles sometimes with yellow spot on dorsal surface. Clypeus smooth and shining. Mesosoma withirregular rugae which are thin in profile. Petiole and postpetiole with a few longitudinal rugae. First gastral tergum weakly punctured
on dorsal surface. Head with sparse pilosity on dorsum. Mesosoma with sparse pilosity on dorsum, hairs of pronotum longer thanpropodeal spines and shorter than that of petiole and postpetiole. Petiole and postpetiole with or without hairs on ventral surface.
Head, mesosoma, petiole and postpetiole varying from yellowish brown to reddish brown, gaster dark reddish brown.
Holotype worker, VIETNAM: Tam Dao, 900 m alt., Vinh Phuc Prov., 9. xi. 1999, VN99-HO-054 (H. Okido) (IEBR).
Paratypes. 3 workers with same data as holotype (KUIC, KUEC).
Other material examined. VIETNAM: Mt. Pia Oac, Bamboo st., 1230 m, Cao Bang Prov. (S. Nomura).
Remarks. M. nomurai belongs to the M. vieti complex. The other members of the complex are: M. guangxiensis and M. vieti sp. nov.
Among those species, M. nomurai can be distinguished from the remains by having the short and sculptured petiole which is expanded
laterally at the anterior portion in dorsal view, and the mesosoma with the longitudinal and parallel rugae.
The species inhabits in soil.
Distribution. VIETNAM: Cao Bang Prov., Vinh Phuc Prov.

Myrmecina padangensis sp. nov.
(Fig. 37)
Holotype worker. TL 3.56, HL 0.88, HW 0.88, CI 100, SL 0.68, SI 77, PW 0.58, ML 0.93.
Paratype worker. TL 3.49, HL 0.86, HW 0.90, CI 104, SL 0.72, SI 80, PW 0.58, ML 0.94 (1 measured).
Worker. Head subrectangular, as long as or shorter than broad, broad posteriorly in full-face view; median portion of occipital margin
concave; occipital corners widely rounded, not projected posteriorly; dorsal outline convex in profile. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a bluntbasal tooth. Dorsal surface of clypeus not concave; median portion of anterior margin slightly projected with or without a median― 80 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 37. Myrmecina padangensis sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face viewof head.

process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired large
denticles, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large andconvex with maximum diameter 0.14 mm, with 8 ommatidia; malar space twice as long as diameter of eye or shorter in profile; distance between occipital margin and posterior margin of eye four times twice as long as diameter of eye or shorter. Antennal scapeshort, just reaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma strongly convex in profile. Pronotum without denticles; anterior portion marginate; anterior ventrolateral portion projected or angulate; anterior dorsolateral portion rounded, not angulate.
― 81 ―

Furrow between pronotum andHirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

mesoepisternal projection variable, narrow to broad. Eumetanotal spine present but small. Propodeal spine triangular, longer thanbroad at base, slightly extending over vertical posteriormost limit of propodeum in profile. Propodeal lobe usually low, sometimes
raised. Propodeal spiracle large, situated near base of propodeal spine, distance between posterior margin of spiracle and posterior
margin of propodeum longer than diameter of spiracle. Petiole long, longer than high in profile, longer than broad in dorsal view;dorsal crest absent; subpetiolar process present, but unclearly without acute anterior apex. Postpetiole slightly broader than petiole indorsal view; anterolateral portion raised; dorsal outline flattened or convex in profile; ventral outline slightly projected with two tops.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae which are thin; ventrolateral portion with longitudinal rugae. Clypeus smooth and shining. Mesosoma
with wavy rugae which are thick. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastralsegment smooth and shining. Head with relatively sparse and long pilosity on dorsum. Mesosoma with sparse and long pilosity on
dorsum, hairs of pronotum as long as propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 2-4 hairs on ventral
surface. Body black to dark reddish brown, forecoxae reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Sumatra, Padang (no collector’s name) (BMNH). Paratypes. 2 workers with same data as holotype(BMNH).
Remarks. M. padangensis does not belong to any species complex. The species can be distinguished from other species by having
the posteriorly broad head, simple anterior margin of the smooth and shining first gastral tergum, the short antennal scape just reaching
occipital corner, the long petiole, and the large eye.
Distribution. INDONESIA: Sumatra.

Myrmecina parallela sp. nov.
(Fig. 38)
Holotype worker. TL 2.98, HL 0.70, HW 0.65, CI 92, SL 0.55, SI 85, PW 0.46, ML 0.78.
Paratype workers. TL 2.82-3.09, HL 0.67-0.72, HW 0.62-0.70, CI 92-97, SL 0.54-0.61, SI 84-95, PW 0.45-0.50, ML 0.74-0.83 (7measured).
Worker. Head subrectangular, slightly longer than broad in full-face view; median portion of occipital margin much concave; occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth);apical tooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave;median portion of anterior margin slightly projected, usually without a median process but sometimes present; lateral portion simple,
lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, close toeach other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes very small circularly and convex,
varying in size with maximum diameter 0.06-0.08 mm and 12-14 ommatidia; malar space twice as long as diameter of eye in profile;
distance between occipital margin and posterior margin of eye five times as long as diameter of eye or longer. Antennal scape long,extending beyond posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion marginate
posteriorly in dorsal view; anterior ventrolateral portion projected. Furrow between pronotum and mesoepisternal projection broad.
Eumetanotal spine present. Propodeal spine elongate, much longer than broad at base, extending over vertical posteriormost limit of― 82 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near base of propodeal spine, apart from margin by its
diameter. Petiole long, longer than high in profile, and longer than broad in dorsal view; dorsal crest located at midlength in profile;subpetiolar process and median longitudinal ridge at ventral portion absent. Postpetiole slightly broader than petiole in dorsal view;dorsal outline flattened or slightly convex in profile; ventral outline slightly projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view.
Head with parallel rugae which are thin and longitudinal; ventrolateral portion variable sculptured, with longitudinal rugae orsmooth and shining. Clypeus smooth and shining. Mesosoma usually with straight rugae longitudinally, sometimes with transverseabcFig. 38. Myrmecina parallela sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

― 83 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

rugae at pronotum. Forecoxa smooth and shining. Petiole and postpetiole with irregular rugae. First gastral segment smooth andshining. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum shorter
than propodeal spine. Petiole with or without 2 hairs on ventral surface. Postpetiole with 4-6 hairs on ventral surface. Head, mesosoma, petiole and postpetiole black, forecoxae and gaster dark reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Sumatra, Padang (no collector’s name) (BMNH). Paratypes. 9 workers with same data as holotype(BMNH).
Remarks. M. parallela does not belong to any species complex. But M. parallela is similar to M. boltoni sp. nov., M. longiseta sp.
nov. and M. monticola sp. nov. on the basis of some characters (see the remarks of M. boltoni). Among those species, M parallela canbe distinguished from the remains by having the anterior clypeal margin without median paired processes and the long propodeal
spine.
Distribution. INDONESIA: Sumatra.

Myrmecina poringensis sp. nov.
(Fig. 39)
Holotype worker. TL 2.45, HL 0.58, HW 0.54, CI 93, SL 0.46, SI 87, PW 0.69, ML 0.61.
Paratype worker. TL 2.34, HL 0.54, HW 0.53, CI 97, SL 0.45, SI 85, PW 0.38, ML 0.59 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (second small tooth or fifth tooth); apical toothstrong, third tooth robust, followed by 4 small teeth, without a basal tooth. Dorsal surface of clypeus not concave; median portion ofanterior margin slightly projected with a median process; lateral portion simple, lacking sharp ridge in front of antennal insertions.
Anterior dorsal surface of labrum with paired small denticles, close to each other. Frontal carinae virtually absent, indistinguishable
from rugae on dorsum of head. Eyes slightly large and moderately convex, varying in size with maximum diameter 0.09-0.10 mm
and 5-6 ommatidia; malar space twice as long as diameter of eye in profile; distance between occipital margin and posterior marginof eye three times as long as diameter of eye. Antennal scape short, just reaching posterolateral corner of head; antennal flange weakly
developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion
marginate; anterior dorsolateral portion angulate; anterior ventrolateral portion not angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine absent. Propodeal spine triangular, shorter than broad at base, clearly not reachingvertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of
propodeum, apart from margin by its diameter. Petiole short, slightly longer than high in profile, longer than broad in dorsal view;dorsal crest located posteriorly in profile; subpetiolar process present with acute anterior apex. Postpetiole much broader than petiole
in dorsal view; anterior portion slightly raised in profile; ventral outline slightly projected rectangularly with two tops.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae which are thin; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma withtransverse rugae on anterior portion, and longitudinal rugae on posterior portion in dorsal view. Forecoxa smooth and shining. Petiole― 84 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

and postpetiole with a few rugae. First gastral segment smooth and shining. Head with relatively sparse and short pilosity on dorsum.
Mesosoma with sparse and long pilosity on dorsum, hairs of pronotum much longer than propodeal spine. Petiole without hairs on
ventral surface. Postpetiole with 2 hairs on ventral surface. Head and mesosoma black or dark reddish brown, petiole, postpetiole andgaster reddish brown, mandibles, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Sabah, Poring Hot Springs, 7. v. 87, 500 m (Burckhardt & Löbl) (BMNH).
Paratypes. 1 worker with same data as holotype; 2 workers with same data as holotype but 10. v. 87, 600 m (BMNH).

abcFig. 39. Myrmecina poringensis sp. nov. a, dorsal view of mesosoma,
petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

― 85 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Remarks. M. poringensis does not belong to any species complex. But M. poringensis is similar to M. dechai sp. nov. on the basisof some characters (see the remarks of M. dechai). Among those species, M. poringensis can be distinguished from the remains by
having the extremely short propodeal spine pointing upward.
Distribution. MALAYSIA: Borneo (Sabah).

Myrmecina raviwonghei Jaitrong, Samung, Waengsothorn & Okido(Fig. 40)Myrmecina raviwonghei Jaitrong, Samung, Waengsothorn & Okido, 2019: 3. Holotype worker (THNHM-I-05470), THAILAND:
Nakhon Ratchasima Province, Wang Nam Kheao District, Udom Sub Subdistrict, 14.46805°N, 101.90416°E, 22.vi.2018 (W. Jaitrong)(WJT220618-6) (THNHM). Paratypes. Twenty-four workers (THNHM-I-05471, MHNG, SKYC, THNHM, USNM), same data as
holotype. [not examined].
Workers. TL 2.50-2.63, HL 0.59-0.62, HW 0.56-0.59, CI 95-96, SL 0.45, SI 76-80, PW 0.40-0.43, ML 0.66-0.69 (2 measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin slightly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth frequently unclear. Dorsal surface ofclypeus concave; median portion of anterior margin projected usually with three processes; lateral portion simple, lacking sharp ridgein front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, which are well separated.
Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes extremely small and moderately convex,
varying in size with maximum diameter 0.05-0.06 mm and 4-6 ommatidia, sometimes 8 ommatidia. Antennal scape short, not reaching posterolateral corner of head; antennal flange developed.
Dorsal outline of mesosoma more or less flattened in profile. Pronotum without denticles; anterior portion marginate; anterior
ventrolateral portion with denticle directing forward and downward. Furrow between pronotum and mesoepisternal projection broad.
Eumetanotal spine present but small. Propodeal spine elongate, directing backward and weakly curved upward, extending over vertical posteriormost limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle medium-sized, situated near posterior marginof propodeum, apart from the margin by its diameter. Petiole short, as long as high in profile, longer than broad in dorsal view; small
but distinct dorsal crest located at midlength in profile; subpetiolar process developed with acute anterior apex. Postpetiole broader
than petiole in dorsal view; dorsal outline flattened or slightly convex in profile; ventral outline distinctly projected with acute anterior
apex.
Anterior margin of gaster concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head puncutred with thin rugae longitudinally; ventrolateral portion punctured with thin rugae. Mandible sometimes with paired
yellow spots on dorsal surface. Clypeus smooth and shining. Mesosoma punctured with thin rugae longitudinally. Forecoxa smoothand shining. Petiole and postpetiole punctured with a few distinct rugae. First gastral tergum punctured dorsally. First gastral
sternum rugulose or punctured. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum,
hairs of pronotum much shorter than propodeal spine. Longest dorsal pilosity of petiole as long as propodeal spine. Petiole without
hairs on ventral surface. Postpetiole with 2-4 hairs on ventral surface. Body black or dark reddish brown, mandibles, forecoxae
reddish brown, antennae and legs yellowish brown.
― 86 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 40. Myrmecina raviwonghei Jaitrong, Samung, Waengsothorn & Okido. a, dorsal
view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole andpostpetiole; c, full-face view of head.

Other material examined. THAILAND: Doi Chiang Dao, 500-600 m alt. nr. Chiang Mai, 19. viii. 1998 (H. Okido) (KUFF): Doi
Chiang Dao, 500-600 m alt. nr. Chiang Mai, 19. viii. 1998 (H. Okido), TH98-HO-015 (KUEC, KUIC); Sakaerat lowland, Necornratchasima (Sk. Yamane).
Remarks. M. raviwonghei belongs to the M. gracilis complex comprising M. curvispina, M. gracilis sp. nov. and M. raviwonghei.
The definition and the distribution of the gracilis complex are given in the latter section. Among those species, M. raviwonghei canbe distinguished from the remains by having the sculptured first gastral sternum without ridge at anterior median portion, and the― 87 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

broad head.
Ergatogynes are present in this species.
Distribution. THAILAND: Chiang Mai Prov., Necornratchasima Prov.

Myrmecina sabahna sp. nov.
(Fig. 41)
Holotype worker. TL 2.86, HL 0.69, HW 0.62, CI 91, SL 0.59, SI 95, PW 0.46, ML 0.74.
Paratype workers. TL 2.88-2.98, HL 0.70-0.74, HW 0.64-0.69, CI 91-93, SL 0.59-0.65, SI 93-94, PW 0.48-0.51, ML 0.69-0.75 (2measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 4 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin slightly projected with four processes, sometimes unclearly; lateral portion simple, lacking sharp ridge in front ofantennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, which are very close to each other.
Frontal carinae present but unclearly, running to about apical two thirds of the line through eyes. Eyes medium-sized and moderately
convex, varying in size with maximum diameter 0.08-0.09 mm and 5-6 ommatidia; malar space twice as long as diameter of eye inprofile; distance between occipital margin and posterior margin of eye four times as long as diameter of eye or longer. Antennal scapeshort, just reaching posterolateral corner of head; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum usually without denticles on dorsolateral portion, but sometimes bearing small denticles; anterior portion marginate but unclearly in dorsal view; anterior dorsolateral portion angulate; anterior
ventrolateral portion variable, not angulate or projected bluntly. Furrow between pronotum and mesoepisternal projection broad.
Eumetanotal spine present but small. Propodeal spine triangular, longer than broad at base, extending over vertical posteriormost
limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, distance
between posterior margin of spiracle and posterior margin of propodeum longer than diameter of spiracle. Petiole short, as long as orslightly longer than high in profile, slightly longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar
process absent. Postpetiole slightly broader than petiole in dorsal view; lateral margin straight; dorsal outline flattened in profile;ventral outline slightly projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with wavy rugae which are thin weakly; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma usually with wavy rugae distinctly. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae. First gastralsegment smooth and shining. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum,
hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with about 6 hairs on ventral
surface. Body black, mandibles, forecoxae reddish brown, antennae and legs yellowish brown.
Holotype worker, MALAYSIA: Mahua Waterfall area, ca. 1000 m alt., Crocker Range N. P., Sabah, 5. xi. 2000, MA00-HO-025 (H.
Okido) (UMS).
Paratypes. 5 workers with same data as holotype (KUEC, KUIC).
― 88 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)abcFig. 41. Myrmecina sabahna sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-faceview of head.

Other material examined. MALAYSIA: East Ridge, Poring Hot Springs, Kinabalu N. P., Sabah (C. Brühl).
Remarks. M. sabahna does not belong to any species complex. But M. sabahna is similar to M. insulana sp. nov. and M. yamanei
sp. nov. on the basis of some characters (see the remarks of M. insulana). The species can be distinguished from the latters by having
the smooth and shining ventrolateral portion of the head, the anterior clypeal margin with median paired processes, and the shortantennal scape just reaching occipital corner.

― 89 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIDistribution. MALAYSIA: Borneo (Sabah).

Myrmecina semipolita Forel
Myrmecina semipolita Forel, 1905: 15. Syntype worker, INDONESIA: Buitenzorg (=Bogor), Java (K. Kraepelin) (depository
unknown) [not seen].
Diagnosis of worker. The following description is based on Forel (1905). Head subrectangular, length shorter than broad in full-faceview. Sides of head slightly convex; median portion of occipital margin straight. Antennal scrobe absent. Mandible smooth andshining with apical and preapical tooth following 4-5 small and unclear teeth. Eyes convex. Antennal scape extending beyond occipital margin of head. Dorsal outline of mesosoma convex without suture. Propodeal spine directing backward, slightly longer thanbroad at base. Eumetanotal spine small. Petiole longer than broad. Postpetiole length shorter than broad, and slightly broader thanpetiole. Gaster oval. Ventrolateral portion of head with rugae. Mesosoma with thick and irregular rugae longitudinally. Remains of
the body smooth and shining. Body black to almost black, antenna, legs, clypeus, mandible reddish.
Remarks. We have not been able to examine type material of M. semipolita.
Distribution. INDONESIA: Java.

Myrmecina spinosa sp. nov.
(Fig. 42)
Holotype worker. TL 4.21, HL 1.06, HW 1.02, CI 97, SL 0.96, SI 94, PW 0.77, ML 1.15.
Paratype workers. TL 3.87-4.10, HL 1.01-1.04, HW 0.99-1.04, CI 97-103, SL 0.83-0.94, SI 84-92, PW 0.74-0.77, ML 1.04-1.12 (3measured).
Worker. Occipital margin strongly concave in full-face view; lateral margin slightly convex; occipital corners projected posteriorly.
Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical tooth strong, third tooth robust, followed
by about 5 small teeth and a stout basal tooth bearing blunt apex. Dorsal surface of clypeus concave; median portion of anterior
margin weakly projected, usually with a distinct median process; lateral portion simple, lacking a sharp ridge, but slightly marginate
on each side, in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles usually distinct, which are well
separated but varied in size and distance between denticles. Frontal carinae virtually absent indistinguishable from rugae on dorsumof head. Eyes medium-sized and convex, varying in size with maximum diameter 0.08-0.14 mm and 6-9 ommatidia. Antennal scapelong, usually extending beyond posterolateral corner of head but length varied; antennal flange developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum with two or four denticles on anterior dorsolateral portion in
dorsal view; anterior ventrolateral portion with denticle directing forward and downward. Eumetanotal spine long with acute apex;
length usually longer than broad at base. Propodeal spine extremely long and curved abruptly upward at their shafts, much extending
over vertical posteriormost limit of propodeum in profile; tips curved outward in dorsal view. Propodeal lobe raised. Propodeal
spiracle situated near base of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum― 90 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

a

d

b

e

c
Fig. 42. Myrmecina spinosa sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, anterior
margin of gaster; e, occipital corner.

slightly longer than diameter of spiracle. Petiole long; small dorsal crest located at midlength in profile; subpetiolar process present;
ventral outline slightly concave. Postpetiole broader than petiole in dorsal view; lateral margin straight and marginated distinctly,
sometimes lateral lobes projected; anterior portion slightly raised in profile; ventral outline projected triangularly.
Anterior margin of gaster concave in dorsal view.
Head weakly punctured with parallel rugae distinctly; ventrolateral portion with longitudinal rugae which are transversely curved
at posterior portion. Mandibles usually with large yellow spots on dorsal surface. Clypeus smooth and shining. Mesosoma withparallel rugae diverging anteriorly (forming “Y”-shape); lateral portion with longitudinal or oblique rugae. Petiole and postpetiole
with a few longitudinal rugae. First gastral tergum smooth and shining. Head and mesosoma with long pilosity on dorsum. Pilosity― 91 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

of pronotum usually longer than broad of apical second segment of antenna. Pilosity of petiole and postpetiole as long as that ofmesosoma. Head, mesosoma, petiole and postpetiole black, legs yellowish, forecoxae varying from reddish brown to black, gasterreddish brown especially posterior portion lighter than anterior.
Holotype worker, MALAYSIA: Mahua Waterfall area, ca. 1000 m alt., Crocker Range N. P., Sabah, 4. xi. 2000, MA00-HO-005 (H.
Okido) (UMS).
Paratypes. 18 workers with same data as holotype; 13 workers with same data as holotype but MA00-HO-006; 3 workers with samedata as holotype but 5. xi. 2000, MA00-HO-026 (KUIC, KUEC).
Other material examined. MALAYSIA: Mahua Waterfall area, ca. 1000 m alt., Crocker Range N. P., Sabah (H. Okido; K. Eguchi;
S. Sonthichai); Sayap Kinabalu, ca. 1000 m alt., Sabah, Borneo (Sk. Yamane); Tawau Hills N. P., Sabah, Borneo (K. Eguchi; Sk.
Yamane); Taman Kinabalu, 1500 m, Sabah, Borneo (Sk. Yamane); Danum Valley, Sabah, Borneo (H. Okido; Sk. Yamane; C. Brühl);Deramakot Forest Reserve, Sabah (C. Brühl); Old Tower R., Lambir N. P., Miri, Sarawak (Sk. Yamane); 50 ha plot, Lambir N. P., Miri,
Sarawak (Sk. Yamane); Sabal, Sarawak, Borneo (Sk. Yamane); Neg. Sembilan, Pasoh For. Res. (M. Brendell, K. Jackson & L. Ficken;
M. Brendell, K. Jackson & S. Lewis). INDONESIA: Sangkimah, Kutai N. P., E. Kalimantan (Sk. Yamane); Mt. Halimon, W-Java (F.
Ito). BRUNEI: Tasek Merimbun (K. Eguchi).
Dimensions of these workers. TL 2.72-3.52, HL 0.72-0.92, HW 0.67-0.85, CI 90-98, SL
0.59-0.86, SI 88-102, PW 0.53-0.62, ML 0.74-0.98 (3 measured).
Remarks. M. spinosa belongs to the M. spinosa complex. The other members of the complex are: M. inflata sp. nov., M. mahuana
sp. nov., M. magnificens and M. tridentata sp. nov. The definition and the distribution of the M. spinosa complex are given in the latter
section. Among those species, M. spinosa can be distinguished from the remains by having the concave anterior margin of the firstgastral tergum, the bent masticatory margin of the mandible, the propodeal spine extending backward, and the head and the mesosoma
with straight and longitudinal rugae. The species shows wide variations in some of characters, such as the lateral lobes of the postpetiole, the position and the shape of denticles on the labrum, the body size (See dimensions), and number and length of the pilosity on
the head and the mesosoma.
The species inhabits in soil and litter.
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah, Sarawak); BRUNEI: Tasek Merimbun; INDONESIA: Borneo (Kalimantan), Java.

Myrmecina striata Emery
Myrmecina striata Emery, 1889: 500. Syntype worker, MYANMAR: Tenasserim, Mt. Mooleyit (L. Fea) (depository unknown) [notseen].
Diagnosis of worker. The following description is based on Emery (1889) and Bingham (1903). Head as long as broad; median
portion of occipital margin concave. Anterior clypeal margin without processes. Antennal scape very nearly extending to occipitalmargin. Dorsal outline of mesosoma slightly convex on dorsum in profile. Propodeal spine directing backward and slightly upward.
Head and mesosoma with longitudinal rugae on dorsal and lateral surface. Petiole and postpetiole with longitudinal rugae on dorsum.
― 92 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Mandible, median portion of clypeus, legs and gaster smooth and shining. Head, mesosoma and gaster with pilosity sparsely. Body
black, mandibles, antennae, legs and gaster reddish yellow.
Remarks. We have not been able to examine type material of M. striata. According to the original descriptions, this species is similar
to M. curvispina and M. raviwonghei.
Distribution. MYANMAR: Tenasserim.

Myrmecina sulawesiana sp. nov.
(Fig. 43)
Holotype worker. TL 2.59, HL 0.61, HW 0.61, CI 100, SL 0.51, SI 84, PW 0.43, ML 0.67.
Paratype workers. TL 2.48-2.61, HL 0.61-0.62, HW 0.59-0.61, CI 96-97, SL 0.50-0.53, SI 84-88, PW 0.40-0.42, ML 0.61-0.69 (3measured).
Worker. Head subrectangular, as long as or slightly longer than broad in full-face view; median portion of occipital margin flattened;
occipital corners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth
tooth); apical tooth strong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave;median portion of anterior margin slightly projected with an unclear median process; lateral portion simple, lacking sharp ridge infront of antennal insertions. Anterior dorsal surface of labrum with paired small denticles, close to each other. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and moderately convex with maximum diameter 0.10 mm,
with 6 ommatidia; malar space twice as long as diameter of eye in profile; distance between occipital margin and posterior margin ofeye three times as long as diameter of eye. Antennal scape short, just reaching posterolateral corner of head; antennal flange weakly
developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion marginate;
anterior ventrolateral portion angulate bluntly. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine
present but small. Propodeal spine triangular, longer than broad at base, just reaching or feebly extending over vertical posteriormost
limit of propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior portion of propodeum, distance
between posterior margin of spiracle and posterior margin of propodeum shorter than diameter of spiracle. Petiole short, slightly
longer than high in profile, slightly longer than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process
present but much reduced, bearing acute anterior apex. Postpetiole broader than petiole in dorsal view; lateral margin straight; dorsal
outline flattened in profile; ventral outline slightly projected with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with wavy rugae which are thin and longitudinal; ventrolateral portion smooth and shining. Clypeus smooth and shining.
Mesosoma with weakly waved rugae longitudinally. Forecoxa smooth and shining. Petiole and postpetiole with longitudinal rugae.
First gastral segment smooth and shining. Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on
dorsum, hairs of pronotum as long as propodeal spine. Petiole and postpetiole without hairs on ventral surface. Body black to reddish
brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Bantimurung, 40 m alt., S. Sulawesi, S 05°01´xE 119°41´, 8. x. 2000, TUS 15 min. #3 (K. Ogata &― 93 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

K. Masaoka) (KUEC).
Paratypes. 4 workers with same data as holotype (KUEC, BZM).
Other materials examined. INDONESIA: Sulawesi, Utara, Dumoga-Bone, N 0°34´xE 123°54´ (M. Horak; no collector’s name)(BMNH); Tengah, Solato R., Taronggo, S 01°45´xE 121°40´ (M. J. D. Brendell) (BMNH); Sulawesi, Utara, G. Muajat, 1780 m (nocollector’s name) (BMNH); N. C. Seram, Manusela N. P., Wae Mual Plain (M. J. D. Brendell) (BMNH).

abcFig. 43. Myrmecina sulawesiana sp. nov. a, dorsal view of mesosoma, petiole andpostpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view ofhead.

― 94 ―

Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Remarks. M. sulawesiana does not belong to any species complex. But M. sulawesiana is similar to M. asiatica sp. nov. on the basisof some characters (see the remarks of M. asiatica). The species can be distinguished from the latter by having the medium-sized andovally eye comprising 15 ommatidia or more.
The species inhabits in litter.
Distribution. INDONESIA: Sulawesi, Seram.

Myrmecina sulcata Emery(Fig. 44)
Myrmecina sulcata Emery, 1887: 449. Syntype workers and male, INDONESIA: Kandari, Celebes (=Sulawesi), iii. 74 (O. Beccari)(MCSN); Buitenzorg (=Bogor) (Ferrari) (depository unknown) [syntypes examined].
Worker. TL 2.57-2.75, HL 0.62-0.66, HW 0.62-0.67, CI 99-104, SL 0.56-0.62, SI 88-93, PW 0.43-0.45, ML 0.62-0.66 (3 measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a basal blunt tooth; small teeth frequently unclear. Dorsal surface of clypeus
usually not concave; median portion of anterior margin projected with a median process but frequently reduced; lateral portion simple,
lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles close to or fused with eachother at base. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes large and convex, varying insize with maximum diameter 0.11-0.13 mm and 7-8 ommatidia; malar space twice as long as diameter of eye or shorter in profile;
distance between occipital margin and posterior margin of eye three times as long as diameter of eye. Antennal scape long, extending
beyond posterolateral corner of head; antennal flange developed.
Dorsal outline of mesosoma convex in profile. Pronotum without denticles; anterior portion not marginate; anterior ventrolateral
portion with denticle, but variable. Furrow between pronotum and mesoepisternal projection more or less broad. Eumetanotal spine
present; the length longer than broad at base. Propodeal spine elongate, extending over vertical posteriormost limit of propodeum in
profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, apart from margin by its diameter.
Petiole long, longer than high in profile, longer than broad in dorsal view; very small dorsal crest located at midlength in profile;subpetiolar process absent, but median longitudinal ridge at ventral portion present. Postpetiole slightly broader than petiole in dorsal
view; lateral margin straight; anterior portion slightly raised in profile; ventral outline not projected.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head with straight rugae distinctly; ventrolateral portion smooth and shining. Clypeus smooth and shining. Mesosoma withstraight rugae distinctly. Forecoxa with transverse rugae, but variable, frequently smooth and shining. Petiole and postpetiole with afew distinct rugae. First gastral segment smooth and shining. Head with sparse and long pilosity on dorsum. Mesosoma with sparseand long pilosity on dorsum, hairs of pronotum slightly longer than apical third segment of antenna. Pilosity of petiole as long as thatof mesosoma. Petiole without hairs on ventral surface. Postpetiole with or without 1-4 hairs on ventral surface. Head and mesosoma
black or dark reddish brown, forecoxae, petiole, postpetiole and gaster reddish brown, mandibles, antennae and legs yellowish brown.
Other material examined. INDONESIA: Wawoteno nr. Kendari, SE Sulawesi (K. Ogata & K. Masaoka) (KUEC).
― 95 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

Remarks. M. sulcata does not belong to any species complex. But M. sulcata is similar to M. muluensis sp. nov. on the basis of somecharacters (see the remarks of M. muluensis). Among those species, M. muluensis sp. nov. can be distinguished from the remains by
having the large eye comprising 20 ommatidia or more, and the broad head.
Distribution. INDONESIA: Sulawesi, Java.

abcFig. 44. Myrmecina sulcata Emery. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Myrmecina sundanica sp. nov.
(Fig. 45)
Holotype worker. TL 2.45, HL 0.56, HW 0.50, CI 89, SL 0.45, SI 90, PW 0.34, ML 0.62.
Paratype worker. TL 2.39, HL 0.54, HW 0.51, CI 94, SL 0.43, SI 84, PW 0.35, ML 0.64 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin concave; occipital corners
rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apical toothstrong, third tooth robust, followed by 5 small teeth and a blunt basal tooth. Dorsal surface of clypeus not concave; median portionof anterior margin feebly projected with a small median process; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrum with paired small denticles bearing acute apex, close to each other. Frontal carinae virtuallyabsent, indistinguishable from rugae on dorsum of head. Eyes extremely small circularly and moderately convex with maximum
diameter 0.06 mm, with 9 ommatidia; malar space twice as long as diameter of eye in profile; distance between occipital margin andposterior margin of eye four times as long as diameter of eye or longer. Antennal scape short, just reaching posterolateral corner ofhead; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles on dorsolateral portion; anterior portion
marginate; anterior ventrolateral portion slightly projected. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present but small. Propodeal spine triangular, longer than broad at base, extending over vertical posteriormost limit ofpropodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of propodeum, apart from marginby its diameter. Petiole short, slightly longer than high in profile, longer than broad in dorsal view; dorsal crest located at midlength
in profile; subpetiolar process present but much reduced, usually bearing acute anterior apex. Postpetiole broader than petiole, 1.5
times as broad as its length except for helcium in dorsal view; anterolateral margin usually rounded; anterior portion slightly raised in
profile; ventral outline projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view; first gastral sternum simple without median longitudinal ridge.
Head weakly punctured with almost staight rugae which are thin; ventrolateral portion usually smooth and shining, sometimes
punctured. Clypeus smooth and shining. Mesosoma weakly punctured with almost straight rugae which are thin and longitudinal.
Forecoxa smooth and shining. Petiole and postpetiole with weak irregular rugae. First gastral tergum punctured dorsally. First gastral
sternum usually smooth and shining, rarely with longitudinal rugae at anterior median portion; lateral portion smooth and shining.
Head with dense and short pilosity on dorsum. Mesosoma with dense and short pilosity on dorsum, hairs of pronotum much shorter
than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 2-4 hairs on ventral surface. Body black to reddish
brown, mandibles, antennae and legs yellowish brown.
Holotype worker, INDONESIA: Kebun Raya, Bogor, W-Java, 15. iv. 1997, FI97-635 (F. Ito) (BZM).
Paratypes. 5 workers and 1 ergatogyne with same data as holotype; 1 worker and 2 ergatogynes with same data as holotype but FI97616; 7 workers and 1 male with same data as holotype but 3. iv. 1997, FI97-545; 2 workers and 1 ergatogyne with same data as holotype but 1. iii. 1997, FI97-38 (KUEC, KUIC).
Other material examined. INDONESIA: W. Bali (IKT. Ginarsa) (BMNH); Java, Bogor (B. Bolton) (BMNH); Botanical Garden,
Bedugul, 1250 m alt., Bali I., S 08°17´xE 115°09´ (J. Abe) (KUEC). MALAYSIA: Ulu Gombak, ca. 250 m alt., Selangor Prov., Malay
Pen. (Sk. Yamane); Sabah, Kibongol Valley, 7 km N. Tambunan (Burckhardt & Löbl) (BMNH).

― 97 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 45. Myrmecina sundanica sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

Remarks. M. sundanica does not belong to any species complex. The species is a distinctive species in having the simple anterior
margin of the sculptured first gastral tergum, and the smooth and shining first gastral sternum. This species corresponds to “Myrmecina sp. F” of Ito et al. (2001). Ergatogynes are present in this species.
The species inhabits in litter.
Distribution. MALAYSIA: The Malay Pen., Borneo (Sabah); INDONESIA: Java, Bali.

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Myrmecina tridentata sp. nov.
(Fig. 46)
Holotype worker. TL 3.34, HL 0.86, HW 0.78, CI 91, SL 0.77, SI 98, PW 0.62, ML 0.93.
Paratype workers. TL 3.28-3.51, HL 0.84-0.94, HW 0.78-0.83, CI 87-93, SL 0.74-0.82, SI 94-100, PW 0.62-0.67, ML 0.91-0.99 (5measured).
Worker. Occipital margin strongly concave in full-face view; occipital corners projected posteriorly. Masticatory margin of mandible
bent at midlength (third small tooth or sixth tooth); apical tooth strong, third tooth acute, followed by 5 small but distinct teeth and astout basal tooth bearing blunt apex. Dorsal surface of clypeus concave; median portion of anterior margin projected, usually withthree distinct processes; lateral portion simple, lacking sharp ridge in front of antennal insertions. Anterior dorsal surface of labrumwith paired small denticles distinctly. Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes mediumsized, varying in size with maximum diameter 0.13-0.14 mm and 7-8 ommatidia. Antennal scape short, just reaching posterolateralcorner of head; antennal flange weakly developed.
Dorsal outline of mesosoma convex in profile. Pronotum with two or four denticles on anterior dorsolateral portion in dorsal
view but frequently unclear; anterior ventrolateral portion with denticle directing forward and downward. Eumetanotal spine triangular distinctly. Propodeal spine elongate, abruptly curved upward at their shafts but varying, distinctly extending over vertical posteriormost limit of propodeum in profile; tips curved outward in dorsal view. Propodeal lobe raised. Propodeal spiracle situated nearbase of propodeal spine, distance between posterior margin of spiracle and posterior margin of propodeum slightly longer than diameter of spiracle. Petiole long; small dorsal crest located at midlength in profile; subpetiolar process triangular; ventral outline slightly
convex or flattened. Postpetiole almost as broad as petiole in dorsal view; lateral margin straight; lateral portion not marginated;dorsal outline slightly convex or almost flattened in profile; ventral outline projected at anterior portion.
Anterior margin of gaster concave in dorsal view;
Head usually with irregular rugae which are wavy, sometimes with longitudinal or parallel rugae; ventrolateral portion marginated on behind eyes in profile, with longitudinal rugae. Mandibles sometimes with small yellow spots on dorsal surface but varying
and unclear. Clypeus smooth and shining. Mesosoma with irregular rugae which are wavy, sometimes with transverse (forming“U”-shape) rugae lateral portion with irregular rugae which are wavy, rarely with longitudinal rugae. Petiole and postpetiole with a
few longitudinal rugae. First gastral tergum smooth and shining. Head with long pilosity on dorsum. Mesosoma with long pilosity
on dorsum, hairs of pronotum usually as long as broad of apical second segment of antenna. Head, mesosoma, petiole and postpetiole
black, legs yellowish brown, forecoxae varying from reddish brown to black, gaster reddish brown.
Holotype worker, MALAYSIA: Sepilok, Sabah, Borneo, 14. viii. 1981 (K. Masuko) (KUEC).
Paratypes. 20 workers and 1 queen (dealate) with same data as holotype (KUIC, KUEC, UMS).
Other material examined. MALAYSIA: Sepilok Forest, Sabah, Borneo (K. Eguchi; Sk. Yamane); Danum Valley, Sabah, Borneo (H.
Okido; Sk. Yamane; C. Brühl); Sayap Kinabalu, Sabah, Borneo (Y. Hashimoto); Tawau Hills N. P., Sabah, Borneo (Sk. Yamane); Mt.
Kinabalu, Sabah, Borneo (Y. Hashimoto); East Ridge, 1530 m, Poring Hot Springs, Kinabalu N. P., Sabah (C. Brühl); Sg., Segerugok,
Song, Sarawak (A. Rahman).
Remarks. M. tridentata belongs to the M. spinosa complex. The other members of the complex are: M. inflata sp. nov., M. mahuana
sp. nov., M. spinosa sp. nov. and M. magnificens. The definition and the distribution of the M. spinosa complex are given in the latter― 99 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 46. Myrmecina tridentata sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view of head.

section. Among those species, M. tridentata can be distinguished from the remains by having the concave anterior margin of the firstgastral tergum, the bent masticatory margin of the mandible, the propodeal spine extending backward, and the head and the mesosoma
with wavy and irregular, or transverse rugae.
The species inhabits in soil.
Distribution. MALAYSIA: Borneo (Sabah, Sarawak).

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Myrmecina undulata Emery
Myrmecina undulata Emery, 1900: 678. Syntype worker and queen, INDONESIA: Si Rambé, Sumatra (E. Modigliani); Pulo Laut(=Laut I.), Borneo (E. Modigliani) (depository unknown) [not seen].
Diagnosis of worker. The following description is based on Emery (1900). Mandible with a few rugae at base. Masticatory marginof mandible with 7-8 acute teeth. Antennal scape long, extending beyond occipital margin. Eumetanotal spine very acute and smaller
than that of M. graminicola (Latreille). Propodeal spine short and triangular. Petiole and postpetiole usual shape, and with longitudinal rugae. Head and mesosoma with wavy rugae. Gaster very shiny. Pilosity long and bristly. Body black and shiny.
Remarks. We have not been able to examine type material of M. undulata.
Distribution. INDONESIA: Sumatra, Laut I. (SE of Borneo).

Myrmecina vieti sp. nov.
(Fig. 47)
Holotype worker. TL 2.45, HL 0.58, HW 0.58, CI 100, SL 0.53, SI 92, PW 0.40, ML 0.64.
Paratype worker. TL 2.61, HL 0.62, HW 0.62, CI 99, SL 0.51, SI 83, PW 0.43, ML 0.72 (1 measured).
Worker. Head subrectangular, as long as broad in full-face view; median portion of occipital margin moderately concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth short, followed by 4-6 small teeth and a stout basal tooth bearing blunt apex. Dorsal surface of clypeus notconcave; median portion of anterior margin weakly projected with blunt and distinct median process; lateral portion raised into a sharp
ridge of shield wall on each side, in front of antennal insertions. Anterior dorsal surface of labrum with paired denticles fused witheach other at base. Frontal carinae present, running back to or beyond level of posterior margin of eyes, but frequently unclear. Eyes
very small, varying in size with maximum diameter 0.05 mm and 5-7 ommatidia. Antennal scape short, just reaching posterolateralcorner of head; antennal flange fully developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior ventrolateral portion more or lessangulate. Eumetanotal spine small but distinct. Propodeal spine triangular, directing upward and backward, not reaching verticalposteriormost limit of the propodeum in profile. Propodeal lobe low. Propodeal spiracle large, situated near posterior margin of
propodeum, apart from margin by its diameter. Petiole longer than high in profile; dorsal crest absent; subpetiolar process projecting
forward with acute apex; ventral outline concave; Postpetiole broader than petiole in dorsal view; lateral margin straight; dorsal
outline flattened in profile; ventral outline projected forward with acute apex at anterior portion in profile.
Anterior margin of gaster concave in dorsal view.
Head with irregular rugae which are thin; ventrolateral portion punctured weakly without rugae. Mandibles smooth and shining,sometimes with yellow spots on anterodorsal surface. Clypeus smooth and shining. Mesosoma with wavy rugae longitudinally.
Petiole and postpetiole almost smooth and shining. Head and mesosoma with sparse pilosity, hairs of head and pronotum as long as
propodeal spines. Pilosity of petiole and postpetiole as long as that of mesosoma; ventral surface with or without hairs. Head, mesosoma, petiole and postpetiole reddish brown, gaster dark reddish brown.
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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

a

b

d

c
Fig. 47. Myrmecina vieti sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b,lateral view of mesosoma, petiole and postpetiole; c, full-face view of head; d, denticles oflabrum.

Holotype worker, VIETNAM: Bavi N. P., 600 m alt., Ha Tai Prov., 12. xi. 1999, VN99-HO-090 (H. Okido) (IEBR).
Paratypes. 4 workers with same data as holotype (KUIC, KUEC); numerous workers and 1 queen (dealate) with same data as holotype
but 670m alt., 21°03´N x 105°22´E, 12. xi. 2001, VN01-HO-111(BZM, KUEC, KUIC, UMS).
Remarks. M. vieti belongs to the M. vieti complex. The other members of the complex are: M. guangxiensis and M. nomurai sp. nov.
Among those species, M. vieti can be distinguished from the remains by having the long and smooth petiole.
The species inhabits under stones.
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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Distribution. VIETNAM: Ha Tai Prov.

Myrmecina yamanei sp. nov.
(Fig. 48)
Holotype worker. TL 2.12, HL 0.51, HW 0.48, CI 94, SL 0.37, SI 77, PW 0.37, ML 0.55.
Paratype worker. TL 2.08, HL 0.49, HW 0.48, CI 98, SL 0.36, SI 75, PW 0.36, ML 0.53 (1 measured).
Worker. Head subrectangular, longer than broad in full-face view; median portion of occipital margin slightly concave; occipitalcorners rounded, not projected posteriorly. Masticatory margin of mandible bent at midlength (third small tooth or sixth tooth); apicaltooth strong, third tooth robust, followed by 5 small teeth and a basal tooth. Dorsal surface of clypeus slightly concave; median
portion of anterior margin feebly projected with a median process or three processes; lateral portion simple, lacking sharp ridge infront of antennal insertions. Anterior dorsal surface of labrum with paired small denticles distinctly, relatively close to each other.
Frontal carinae virtually absent, indistinguishable from rugae on dorsum of head. Eyes small ovally, and moderately convex, varying
in size with maximum diameter 0.06 mm and 12-14 ommatidia; malar space twice as long as diameter of eye or longer in profile;
distance between occipital margin and posterior margin of eye four times as long as diameter of eye or longer. Antennal scape short,
not reaching posterolateral corner of head; antennal flange weakly developed.
Dorsal outline of mesosoma slightly convex in profile. Pronotum without denticles; anterior portion marginate; anterior ventrolateral portion angulate. Furrow between pronotum and mesoepisternal projection broad. Eumetanotal spine present but small.
Propodeal spine triangular, just reaching or slightly extending over vertical posteriormost limit of propodeum in profile. Propodeal
lobe low. Propodeal spiracle medium-sized to large, situated near posterior margin of propodeum, distance between posterior marginof spiracle and posterior margin of propodeum variable. Petiole short, as long as or slightly longer than high in profile, slightly longer
than broad in dorsal view; dorsal crest located at midlength in profile; subpetiolar process variously developed from weakly raised
median longitudinal ridge to distinct projection. Postpetiole broader than petiole, 1.5 times longer than its length except for helcium
in dorsal view; dorsal outline flattened or slightly convex in profile; ventral outline projected rectangularly with acute anterior apex.
Anterior margin of gaster not concave in dorsal view.
Head punctured with straight rugae diverging posteriorly; ventrolateral portion sculptured variable, with or without irregularrugae. Clypeus smooth and shining. Mesosoma punctured with parallel rugae which are thick and longitudinal. Forecoxa smoothand shining. Petiole and postpetiole with longitudinal rugae. First gastral segment smooth and shining. Head with more or less dense
and short pilosity on dorsum. Mesosoma more or less with dense and short pilosity on dorsum, hairs of pronotum shorter than propodeal spine. Petiole without hairs on ventral surface. Postpetiole with 2-4 hairs on ventral surface. Body black, forecoxae black todark reddish brown, mandibles, antennae and legs brown to yellowish brown.
Holotype worker, MALAYSIA: Ulu Gombak, ca. 250 m alt., Selangor Prov., Malay Pen., 5. vii. 1999 (Sk. Yamane) (UMS).
Paratypes. 1 worker with same data as holotype; MALAYSIA: Neg. Sembilan, Pasoh For. Res., xi. 1994 (M. Brendell, K. Jackson &S. Lewis); Malaya, Selangor, Gombak, 15. x. 84 (C. Betts) (UMS, KUEC, KUIC).
Remarks. M. yamanei does not belong to any species complex. But the species is similar to M. insulana sp. nov. and M. sabahna sp.
nov. on the basis of some characters (see the remarks of M. insulana). The species can be distinguished from the latters by having theshort antennal scape not reaching occipital corner by the length of scape width (SI≤77), and the anterior clypeal margin with a median― 103 ―

Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHIabc
Fig. 48. Myrmecina yamanei sp. nov. a, dorsal view of mesosoma, petiole and postpetiole; b, lateral view of mesosoma, petiole and postpetiole; c, full-face view ofhead.

process.
The species inhabits in litter.
Distribution. MALAYSIA: The Malay Pen.

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Taxonomic revision of the ant genus Myrmecina in Southeast Asia (Hymenoptera: Formicidae)

Acknowledgements
We would like to thank Dr. Hiroshi Shima (Kyushu Univ., Japan) for his continuing guidance and advising, Dr. Seiki Yamane (KUIC)
for his kindly support and advising, Dr. Fuminori Ito (Kagawa Univ., Japan) for his providing with huge and precious specimens, andDr. Osamu Yata and Dr. Kunio Araya (Kyushu Univ., Japan) for their advising. I also wish to thank Mr. Barry Bolton & Ms. Christine
Taylor (BMNH), Dr. Carsten Brühl (Biocenter, Germany), Dr. Daniel Burkhardt (Naturhistorisches Museum, Switzerland), Dr. James
Carpenter (AMNH), Dr. Katsuyuki Eguchi (KUIC), Mr. Toru Kikuta, Dr. Chung-Chi Lin (NTU), Dr. Bernhard Merz (MHNG), Dr.
Maryati bte Mohamed (UMS), Dr. Shûhei Nomura (NSMT), Dr. Kyohsuke Ohkarawa (Kanazawa Univ., Japan), Dr. Valter Raineri(MCSN), Dr. Ted R Schultz & Mr. David G. Furth (USNM), Dr. Stefan Schödl (NHMW), Dr. Mamoru Terayama, Dr. Kyoichiro Ueda
(Kitakyushu Museum & Institute of Natural History, Japan), Dr. Bui Tuan Viet, (IEBR), Dr. Philip S. Ward (University of California,
U. S. A), Mr. Decha Wiwatwitaya (KUFF), Dr. Lajos Zombori (HNHM) for their lending of type materials and precious specimens,or providing precious specimens. Finally we would like to appreciate the ANeT members and all my friends for their hospitality andencouragement.
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Sociobiology, 52: 283-291.

Received February 15, 2020; accepted March 15, 2020

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Hirofumi OKIDO, Kazuo OGATA & Shingo HOSOISHI

東南アジアにおけるカドフシアリ属の分類学的再検討（ハチ目：アリ科）
大城戸博文1）・緒方

一夫2）・細石真吾2）

一般財団法人九州環境管理協会公益活動推進センター
2）
九州大学熱帯農学研究センター

1）

要旨：東南アジア産カドフシアリ属 Myrmecina（アリ科フタフシアリ亜科）について分類学的再検討を行い、40 新種を含む53種を整理した。このうち16種について４種群を設立した。働きアリを用いた種の検索表を作成した。

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