Chapter 9 IDENTIFICATION GUIDE TO THE ANT

GENERA OF BORNEO
Yoshiaki HASHIMOTO

Introduction
Ants are one of the most abundant and diverse animal groups in tropical ecosystems (Stork, 1987, 1991),
and they function at many levels in these ecosystems - as predators and prey, as detritivores, mutualists, and
herbivores (Hölldobler and Wilson, 1990). Thus, ants have the potential to yield more meaningful biodiversity
data than many other organisms, such as plants, birds, and butterflies. Moreover, since most species have
stationary, perennial nests with fairly restricted foraging ranges, ants have a potential role as indicators of
environmental change. Because of the potential usefulness, inventory of ants has been viewed as an important
task in tropical biodiversity and conservation studies (Agosti et al., 2000).

The most difficult part of ant inventory in tropical region is identification process. Inventory data are usually
analyzed by relying on the presence or absence of species. However, identification of tropical ant specimens
to species will be very difficult or impossible, because most groups of the ants have yet to be studied in
detail. This difficulty makes the recognition of morphospecies a necessary part of inventory studies for ants
(Agosti et al., 2000).

The identifying ants to genus-level are not impossible, because excellent identification-key to ant genera of
the all parts of the world is available in Bolton (1994). Thus, for sorting ant specimens into morphospecies,
they should be identified to genus (i.e., “ Ant species 1 and species 2” to “Aenictus sp. 1 and Camponotus
sp.1”). This makes it easy to handle and analyze the data. Furthermore, ecological information, such as food
habits, nest-site preference, colony size, etc., is provided to many genera (Hölldobler and Wilson, 1990).
Thus identifying ants to genus-level can provide useful information on environmental monitoring,
conservation evaluation, and ecological research.

However, the key of Bolton (1994) is technical, and requires some knowledge of taxonomic descriptions.
For no-taxonomist, a more user-friendly and pictorial identification key to ant genera is need. Furthermore,
for local inventory the regional identification key may be more convenient, because only a subset of the
genera is found in each geographic region. In this chapter, therefore, I provide a pictorial identification key
to Bornean ant genera.

Before attempting to identify an ant specimen, knowledge of the specimen mounting technique and the
external anatomy must be confirmed. The chapter also provides instruction of the mounting techniques and
glossary of ant morphology terms.

Outline of ant diversity in Borneo

The ant fauna of Borneo Island is very diverse and unique. The island may have 9 subfamilies, 94 genera and
more than 1000 described species. Worldwide, there are 16 subfamilies, about 300 genera and about 15,000
described species of ants (Bolton 1995). Thus Borneo may have representatives of about 30% of its genera
and about 7% of its species, though Borneo covers less than 0.2% of the earth’s land surface. The 6 genera,
including Bregmatomyrma, Epelysidris, Ishakidris, Loweriella, Secostruma, Tetheamyrma, may be endemic to Borneo.
In Indo-Australian region, which include Malaysia, Philippines, Indonesia, New Guinea and Pacific Ocean
islands, 22 endemic genera are found (Bolton, 1995). Thus, about 27% of them are Bornean ants.

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Borneo has tropical rain forests with the richest plant diversity in the world. This is one of reason why the
island has very diverse and unique ant fauna. Ant fauna in Borneo, especially in the canopy of tropical rain
forests, has yet to be studied in detail. As the ant inventory are completed, many additional or new genera
and species should be found from Borneo.

Notes on the keys

The keys provided here are designed to identify workers only. This is because workers are the most
commonly encountered caste of ants in inventory. For sake of no-ant specialist user, I try to select and use
easily observable characters in the keys. However, the use of a microscope is essential for identification of
ants, because of their small size.

To make reliable identification, it is advisable to check additional information to confirm your

identification. The additional information includes the diagnostic characters and biological information of
taxonomic group, and the known distribution of the group. The book of Hölldobler and Wilson (1990),
and Social insect Web-site (http://research.amnh.org/entomology/social_insects/) can help you to search
such information. This chapter also provide genus list of Bornean ants, with their taxonomic references and
biological data.

In some case specimen may not identified certainly. When this happens, you try to use Bolton’s key (1994).
The illustrations provided here should help you to understand the taxonomic descriptions in the key.

Preparation and Preservation of ant specimens Fig. 1

The preparation of ant specimen is more important than
the collection of specimen in the field. Big size organisms,
like mammal and birds, can be identified in the field.
However, small organisms, like ants, are needed to preserve
and prepare for identification in the laboratory. The use
of good preservation and preparation techniques serves
to facilitate the identification of species and improves
research value of specimen. Preparation and preservation
techniques of ant specimens, here, are outlined.

1) Pinning Specimens
Commonly used specimen mounting tools
Insect specimens are mounted on pins so that they may include a pinning block, forcep, pins, points, glue.
be handled and examined with the greatest convenience.
Therefore, for identification and taxonomic study, pinning specimens should be used. Fig. 1 shows typical
tools needed to make pinning specimens.

Ants are usually too small to be pinned directly, and so should be pinned as double mounts, i.e., the
specimen is glued to a card point and the point is pinned through the broad end with a No. 3 insect pin
(Fig. 2). Card points are slender little triangles of stiff paper, which are cut from a strip of paper (they should
be no more than 10 mm long and 5 mm wide). The choice of good quality paper for card points is important
to prevent the specimen from working lose and rotating on the pin. To mount the specimens on card points,
white glue is used commonly. Touch the tip of the point to the glue, and then the point is attached to the

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Fig. 2 - Pinning ant specimen

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platform formed by the middle and hind coxae, inserting the point from the right side. Only a small amount
of glue should be used, since excessive glue may obscure certain structures necessary for identification. The
height of the card point on the pin will depend somewhat on specimen size, but enough of the pin should
always be exposed above it to be grasped without the fingers touching and possibly damaging the specimen.
Good height may be obtained by using the pinning block.

Fig. 3 illustrates some right and wrong examples for mounting specimen. In well-mounted specimen, the
side of the alitrunk, the dorsal profile, and the dorsal and ventral margins of the petiole are clearly visible
(Fig. 3A). Otherwise, in poorly-mounted specimen, the legs are projecting upward, obscuring the dorsal
profile and the petiole (Fig. 3B). In ants, the area around the alitrunk and petiole bear many identification
characters, and therefore the parts of the body should be free for easy examination. The head is also important
area for identification. It is advisable to push upward a pair of antennae, so that they do not obscure the
head (Fig. 3C). In many genera, the palp formula and mandibular dentition are diagnostic characters. One
or more specimens in a series should have the mandibles opened and the mouthparts everted prior to
mounting. It is somewhat troublesome and time-consuming work to make good mounted specimens.
However, if any identification key is to be used with some hope of success, specimens must be mounted in
the way that the characters needed for identification are clearly visible.

A: Well-mounted specimen.
B: Poorly-mounted specimen.
C: Desirable position of antennae and mandibles.

2) Labeling
Specimens without data labels have no scientific value. Therefore, during preparation and mounting,
specimens should bear temporary data labels, and any time a sample is subdivided, the label must be copied
so that every specimen continues to be accompanied by the data. Here, the important points for making the
labels are described.

A) Paper and Size of Label

The paper used for making labels should be thick enough so that the labels remain flat and do not rotate
loosely on the pin. Label size of insect specimen may depend on the size of the insect on a pin. An advantage
of a label that exceeds the size of the insect is that if the specimen is accidentally dropped, the label may keep
the insect from being damaged. However, for ant specimens, it is proper that the size should be no more
than 12 mm long and 3 mm wide (Fig. 2), because most ants are very small and large labels are inconvenient
to handle and examine specimen. If more data are included, more than one label should be used.

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B) Label printing
The style and technique of label printing or writing may vary from one worker to another. Recently,
computer-generated labels printed by laser printers become increasingly common. This way has made it
easier to produce labels in very small font sizes. And word processor and database software that help in
generating this kind of label is widely available. However, toner of laser printer may deteriorate with age so
that laser printed label will peeled off the toner finally. Methods of computer-generated labels are subject to
improvement. The best labels may still be professionally printed labels.

C) Label Data
Collection locality, date (day, month and year), and collector name is indispensable specimen data. These
data should be printed on a label (Fig. 2). Furthermore, since Global Positioning System (GPS) is available
now, it is recommendable to put latitude and longitude on a label in addition to the primary locality data.
For ants, information about collecting or nesting site, such as soil, leaf-letter and tree, are of great importance
for identification purposes, and so usually recorded on additional labels. When a colony can be collected, it
is advisable that colony code-number is assigned to the specimens belonged to the colony. This way ensures
to refer combination with different castes from same colony.

D) Placing the Labels

The pin is inserted through the center of the right side of the label (Fig. 2), with the long axis of the label
oriented in the same direction as the card point.

Wet specimen - Liquid Preservation

In ants, liquid preservation is well used for duplication specimen storage,
and also temporary storage of ant specimens, until the specimen can be
mounted. Most commonly, 70-80 % ethanol is used as preservation fluids.
Fig. 4 shows typical tools needed to preserve wet-specimens. The
specimens are kept in a small vial and then the vials are kept in a jar. Each
vial should be individually labeled with complete collection data. Labels
may also be placed on the outside of the jars to indicate the enclosed
contents. Special care should be taken with labels for wet-specimens.
Typewritten labels and laser printed labels are generally unacceptable,
since such labels cannot withstand the constant exposure to the alcohol.
The best may be the labels writing with soft lead pencil or India inks.
Each vial and jar should be filled with alcohol to the top. And, the jars
should be checked periodically to prevent alcohol evaporation.

Preservation for Molecular and Chemotaxonomic studies

In ants, specimens for molecular work should be collected in absolute
(100%) ethyl alcohol. It is best that specimens are thoroughly dehydrated
by changing the alcohol at least a couple of time before the specimens
are stored for any length of time. It is also advisable to keep the specimens
in refrigerator. For chemotaxonomic analysis of body-surface wax, ant Wet specimen.
specimen should be kept in less than 70% ethyl alcohol, to prevent
dissolution of the wax into preservation fluid.

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References
- Agosti, D., J. Majer, L. Alonso, and T. R. Schultz (eds.) (2000) Ants: Standard methods for measuring and
monitoring biodiversity. 280 pp. Smithsonian Institution Press.

- Bolton, B.0(1994) Identification guide to the ant genera of the world. Harvard University Press, Cambridge, MA.
222 pp.

- Bolton, B. (1995) A new general catalogue of the ants of the world. Harvard University Press, Cambridge, MA.
504 pp.

- Hölldobler, B. and Wilson, E. O. (1990) The Ants. 733 pp. Belknap Press of Harvard University, Cambridge,
MA.

- Stork, N. E. (1987) Guild structure of arthropod fauna of Bornean lowland rain forest trees. Ecol. Entomol.,
12: 69-80

- Stork, N. E. (1991) The composition of arthropod fauna of Bornean lowland rain forest trees. J. Trop. Ecol. 7:161-180

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KEY TO BORNEAN SUBFAMILY OF FORMICIDAE

(No. 1)

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KEY TO BORNEAN SUBFAMILY OF FORMICIDAE

(No. 2)

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KEY TO BORNEAN SUBFAMILY OF FORMICIDAE

(No. 3)

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KEY TO BORNEAN SUBFAMILY OF FORMICIDAE

(No. 4)

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Key to Bornean Genera of the Subfamily Dolichodeinae

(No. 1)

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Key to Bornean Genera of the Subfamily Dolichodeinae

(No. 2)

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Key to Bornean Genera of the Subfamily Dolichodeinae

(No. 3)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 1)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 2)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 3)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 3-1)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 4)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 4-1)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 4-2)

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Key to Bornean Genera of the Subfamily Formicinae

(No. 5)

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Key to Bornean Genera of the Subfamily Leptanillinae

(No. 1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 2)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 3)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 4)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 5)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 5-1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 6)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 6-1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 6-2)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 6-2-1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 6-2-2)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7-1)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7-2)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7-3)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7-4)

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Key to Bornean Genera of the Subfamily Myrmicinae

(No. 7-5)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 1)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 2)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 3)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-1)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-2)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-3)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-4)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-5)

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Key to Bornean Genera of the Subfamily Ponerinae

(No. 4-6)

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Glossary of Ant Morphology

Abdomen Acidopore (AC)
The abdomen in worker ants consists of seven It is the orifice of the formic acid projecting system,
visible segments (A1-7). The first abdominal which is formed from apex of the hypopygium,
segment is the propodeum (PPD, A1), which appearing a short nozzle, generally with a fringe of
immovably fused to the thorax. The second short setae.
abdominal segment is the petiole (PT, A2).
Abdominal segment 3 is the first gastral segment
when it is full-sized and broadly articulated to the
following segment (A3), but when reduced and
isolated it is called the postpetiole (PPT).
Abdominal segment 3 or 4 through to 7 is called
the gaster (GA). The last visible abdominal tergite
(A7) is the pygidium (PY), and the last visible
sternite is the hypopygium (HY).
See Petiole.

Alitrunk (Mesosoma)
The alitrunk consists of the three segments of the
true thorax (pro-, meso-, and metathorax) to which
is fused the propodeum (the tergite of the first
abdominal segment), to form a single unit (AL).

Pronotum (PN): The dorsal sclerite of the prothorax.

In ants, the pronotum extends across to dorsum and
down the sides of the prothorax.

Propleuron (PR): The lateral part of the prothorax.

In ants, the propleuron is concealed by the lateral
part of the propleuron.

Mesonotum (MS): The dorsal part of the mescthorax.

In ants, the mesonotum may be separated from the
pronotum by the promesonotal suture (PMS), or
may be fused to it to form a single sclerite, the
promesonotum (prs).

Mesopleuron (MSP): The lateral and ventral part of

the mesothorax. The mesopleuron may consist of a
single sclerite or may be divided by a transverse
groove into an upper and a lower part .

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Metanotum (MTN): The dorsal sclerite of the of three parts, scape (SC), pedicel (PD) and
metathorax. In ants, the metanotum may be funiculus (= flagellum, FU). The antenna in ants
reduced, or obliterated. consists of 4-12 segments. The funicular segments
may be filiform or enlarged to form a club.
Metanotal groove (MTG): In ants, the mesonotum
and propodeum are often separated by a transverse
groove or impression (Metanotal groove)
representing the last vestige of the metanotum.

Metapleuron (MTP): The lateral and ventral part of

the metathorax. In ants, the metapleuron is located
posteriorly on the side of the alitrunk, below the
level of the propodeum. The metapleuron bears
metapleural grand (MG).
See metapleural grand
Atennal scrobe (ASC)
It is a groove, impression, or excavation in the side
of the head, which runs above or below the eye, for
reception of antennal scapes.

Antennal socket (insertion)

The scape articulates with the head in the antennal
socket (ASO). In ants, the antennal socets usually
are overhung and concealed the fronta lobe, but in
some the fronta lobe absent or reduce so that they
are exposed.

Antenna
It is a paired segmented sensory appendage of the
head between the compound eyes, which consists

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Club (CL) Frontal carina (FC)

The enlarged apical funicular segments of an A pair of longitudinal ridges on the head, located
antenna. In ants, apical 1-4 segments may be dorsally behind the clypeus and between the
enlarged to form a club. antennal sockets. Commonly the frontal carinae
anteriorly are expanded into the frontal lobes.
See Frontal lobes

Clypeus (CP)
Anterior sclerite of the dorsal head, bounded Frontal lobe (FB)
posteriorly by the fronto-clypeal suture (= posterior The frontal lobes commonly are extensions, which
clypeal margin or border, FS). The anterior clypeal partially or entirely cover and conceal the antennal
margin usually forms the anterior margin of the sockets.
head in full-face view. The body of the clypeus
consists of median portion and a pair of lateral
portions (MC, LC). The median portion of the
clypeus may be equipped with one or more
longitudinal carinae, or may be variously specialized
in shape.

In some taxa the clypeus is very reduced and

extremely narrow from front to back.

Leg segments
Each leg consists of a basal coxa (BC) that articulates
with the alitrunk, followed in order by a trochanter
(TR), femur (FE), a tibia (TB), and a tarsus (TA), the
last consisting of five small segments and
terminating apically in a pair of claws (CA).

Tibial spur (TBS) A socketed spur located at the apex

of each tibia. The forelegs have a single pectinate
tibial spur, modified into an antennal cleaner
(strigil). The middle and hind legs may each have
two, one, or no spurs present.

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Teeth (T): Teeth are usually sharp and triangular in

shape but may be rounded (crenulate), long, narrow,
and spine-like (spinlform), or peg-like. The tooth
at or nearest to the basal angle is the basal tooth
(BT), and the distalmost tooth is apical tooth (AT).
When teeth or a combination of teeth and denticles
are present, the mandible is dentate. If only tiny
denticles occur the mandible is denticulate, and if
the margin lacks teeth and denticles it is edentate.

Denticles (DN): Denticles are short or very educed

acute teeth.

Diastema (DI): A diastema is a gap in a row of teeth.

Mandible (MD)
The paired, heavily sclerotized appendage of the
mouthparts between the labrum and maxilla (See
Mouth parts). In ants, the mandibular margins
usually form a triangular or subtriangular shape in
full-face view, but in some become from elongate-
triangular to linear mandibles. Extremely curved
mandibles, usually quite short and with few or no
teeth are termed falcate.

Apical margin (=masticatory margin, MA): In

full-face view, with the mandibles closed, the apical
margin forms inner margin or border of each
mandibular blade, and is usually armed with teeth.

Basal margin (MB): The basal margin of each

mandible is transverse or oblique margin closest to
an anterior clypeal margin. The apical and basal
margins may join through a curve, or meet in an
angle (Basal angle, BA). When the mandibles are
narrow or linear, the basal angle may be lost. In a
few taxa teeth may occur on the basal margin, but
in most this margin is unarmed.

External margin (=lateral margin, ME): The

external margin of each mandible forms its outer
border.

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Y. HASHIMOTO

Metapleural grand (MG)

The metapleural grand is an exocrine gland whose
orifice is usually situated in the posteroventral
corner of the side of the alitrunk, above the level of
the metacoxa and below the level of the propodeal
spiracle.

Mouthparts
The appendages of the head used for feeding, Occipital carina (OC)
including the labrum, hypopharynx, mandibles, A ridge on the posterior surface of the head that
maxillae, and labium. separates the occiput from the vertex and gena.

Maxillary palps (MP): The segmented sensory palps

of the maxillae. Each palp may have at most 6
segments but these are variously reduced in number
in different ant groups.

Labial palps (LP): The segmented sensory palps of

the abium. Each palp may have at most 4 segments
but these are variously reduced in number in Petiole (PT)
different ant groups. A way of indicating the number The second abdominal segment (see Abdomen). The
of segment in the maxillary and labial palps is petiole takes the form of a node (nodiform) or of a
termed as Palp Formula (PF). The number of scale (squamiform), but in some taxa it may be
maxillary palp segments is given first, the number represented by only a narrow, subcylindrical
of labial palp segments second. segment.

Labrum (LB): Mouthpart sclerite that hinges on the Peduncle (PD): The relatively narrow anterior section
anterior margin of the clypeus and usually folds of the petiole. when the peduncle is present, the
back and down over the apices of the maxillae and petiole is termed pedunculate. When the peduncle
labium when the mouthparts are not in use. is absent, so that the node or scale of the petiole
immediately follows the articulation with
propodeum, the petiole is termed sessile.

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Spongiform (SP)
Specialized sponge-like external cuticular tissue,
distributed mainly about the waist segments in some
groups of ants.

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Y. HASHIMOTO

Genus list of Bornean Ants

Subfamily Aenictinae
Tribe Anenictini
Genus Aenictus Shuckard 1840
Type species: Aenictus ambiguous, by original designation.
Taxonomic Reference
Shuckard, W. E. 1840. Monograph of the Dorylidae, a family of the Hymenoptera Heterogyna. Ann. Mag. Nat.
Hist. 5: 258-271.
Wilson, E.O. 1964 The true army ants of the Indo-Australian area. Pacific Insects 6: 427-483
Biology: Army ants (Predators of other ants)

Subfamily Cerapachyinae
Tribe Cerapachyini
Genus Cerapachys Smith 1857
Type species: Cerapachys antennatus, designated by Bingham (1903).
Taxonomic References
Smith, F. 1857. Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca;
and at Singapore, by A. R. Wallace. J. Proc. Linn. Soc. London Zool. 2: 42-88.
Brown, W.L.,Jr. 1975 Contributions toward a reclassification of the Formicidae. 5. Ponerinae, tribes Platythyreini,
Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini. Search Agriculture 5: Entomology (Ithaca)
15 : 1-115
Biology: Army ants (Predators of other ants)

Subfamily Dolichoderinae
Tribe Dolichoderini
Genus Bothriomyrmex Emery 1869
Type species: Bothriomyrmex costae Emery, by monotypy.
Taxonomic Reference
Emery, C. 1869. Descrizione di una nuova formica italiana. Ann. Mus. Zool. R. Univ. Napoli 5: 117-118.
Shattuck, S.O. 1992 Generic revision of the ant subfamily Dolichoderinae. Sociobiogy 21: 1-181
Habitat: Wide variety of habitats, from grasslands to lowland rain forests. Nests in soil or in rotten wood
Biology: Foraging on ground and trees. Some are temporary sociall parasites of Tapinoma and Iridomyrmex.

Genus Dolichoderus Lund 1831

Type species: Formica attelaboides, by monotypy.
Taxonomic References
Lund, P. W. 1831. Lettre sur les habitudes de quelques fourmis du Bresil, adressee a M. Audouin. Ann. Sci. Nat.
23: 113-138.
Shattuck, S.O. 1992 Generic revision of the ant subfamily Dolichoderinae. Sociobiogy 21: 1-181
Habitat: Arboreal
Biology: Generalized foragers, tend homopteras

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Genus Iridomyrmex Mayr 1862

Type species: Formica detecta, designated by Bingham (1903).
Taxonomic Reference
Mayr, G. 1862. Myrmecologische Studien. Verh. Zool.-Bot. Ges. Wien 12: 649-776.
Shattuck, S.O. 1992 Generic revision of the ant subfamily Dolichoderinae. Sociobiogy 21: 1-181
Habitat: Wide variety of habitats, from grasslands to lowland rain forests. Nests in soil or in rotten wood
Biology: Generalized foragers, tend homopteras

Genus Loweriella Shattuck 1992

Type species: Loweriella boltoni, by original designation.
Taxonomic References
Shattuck, S. O. 1992. Generic revision of the ant subfamily Dolichoderinae (Hymenoptera: Formicidae).
Sociobiology 21: 1-181.
Habitat: Leaf litter in rain forests
Note: The single species, Loweriella boltoni, is known from Sarawak and Brunei

Genus Philidris Shattuck 1992

Type species: Formica cordata, by original designation.
Taxonomic References
Shattuck, S. O. 1992. Review of the dolichoderine ant genus Iridomyrmex Mayr with descriptions of three new
genera (Hymenoptera: Formicidae). J. Aust. Entomol. Soc. 31: 13-18.
Habitat: Forests
Biology: Generalized foragers, tend homopteras

Genus Tapinoma Foerster 1850

Type species: Tapinoma collina, by monotypy.
Taxonomic Reference
Foerster, A.. 1850. Hymenopterologische Studien. 1. Formicariae. , Aachen. 74 pp.
Shattuck, S.O. 1992 Generic revision of the ant subfamily Dolichoderinae. Sociobiogy 21: 1-181
Habitat: Wide variety of habitats, from grasslands to lowland rain forests. Nests in soil or in rotten wood
Biology: Generalized foragers, tend homopteras
Note: One species, T. melanocephalum, is a tramp species widely distributed in tropical regions

Genus Technomyrmex Mayr 1872

Type species: Technomyrmex strenuus, by monotypy.
Taxonomic Reference
Mayr, G. 1872. Formicidae Borneenses collectae a J. Doria et O. Beccari in territorio Sarawak annis 1865-1867.
Ann. Mus. Civ. Stor. Nat. Genova 2: 133-155.
Shattuck, S.O. 1992 Generic revision of the ant subfamily Dolichoderinae. Sociobiogy 21: 1 181
Habitat: Moist forests. Nests in soil, in branches, or in carton nests under leaves or tree trunks.
Biology: Generalized foragers, tend homopteras
Note: One species, T. albipes, is a tramp species widely distributed in tropical regions

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Y. HASHIMOTO

Subfamily Dorylinae
Tribe Dorylini
Genus Dorylus Fabricius 1793
Type species: Vespa helvola, by monotypy.
Taxonomic Reference
Fabricius, J. C.. 1793. Entomologia systematica emendata et aucta. Vol. 2. Christ. Gottl. Proft, Hafniae. 519 pp.
Wilson, E.O. 1964 The true army ants of the Indo-Australian area. Pacific Insects 6: 427-483
Biology: Army ants. Generalized predators

Subfamily Formicinae
Tribe Brachymyrmecini
Genus Cladomyrma Wheeler 1920
Type species: Aphomomyrmex hewitti, by original designation.
Taxonomic Reference
Wheeler, W. M. 1920 The subfamilies of Formicidae, and other taxonomic notes. Psyche 27: 46-55.
Agosti, D. 1991 Revision of the Oriental ant genus Cladomyrma, with an out-line of the higher classification of the
Formicinae Systematic Entomology 16: 293-310
Habitat: Nests in live pithy
Biology: Generalized foragers. Tend coccoids (Pseudococcidae) inside the nest hollows.

Tribe Bregmatomyrmini
Genus Bregmatomyrma Wheeler, 1929
Type species: Bregmatomyrma carnosa Wheeler, by original designation.
Taxonomic Reference
Wheeler, W. M. 1929. Three new genera of ants from the Dutch East Indies. Amer. Mus. Nov. 349: 1-8.
Note: Only single species, B. carnosa, (queen only) is known from Borneo.

Tribe Camponotini
Genus Camponotus Mayr 1861
Type species: Formica ligniperda Latreille, designated by Bingham (1903).
Taxonomic Reference
Mayr, G. 1861 Die Europ‰ischen Formiciden. (Ameisen.)., Vienna. 80 pp.
Habitat: Nests in ground, in dead wood, or in trees.
Biology: Generalized foragers.

Genus Echinopla Smith 1857

Type species: Echinopla melanarctos, designated by Wheeler (1911).
Taxonomic Reference
Smith, F. 1857 Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca;
and at Singapore, by A. R. Wallace. J. Proc. Linn. Soc. London Zool. 2: 42-88.
Habitat: Arboreal
Biology: Foraging on trees

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Genus Polyrhachis Smith 1857

Type species: Formica bihamata, by original designation.
Taxonomic Reference
Smith, F. 1857 Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca;
and at Singapore, by A. R. Wallace. J. Proc. Linn. Soc. London Zool. 2: 42-88.
Habitat: Many arboreal. some in silk-woven leaf nests.
Biology: Generalized foragers

Tribe Gesomyrmecini
Genus Gesomyrmex Mayr 1868
Type species: Gesomyrmex hoernesi, by monotypy.
Taxonomic Reference
Mayr, G. 1868 Die Ameisen des baltischen Bernsteins. Beitr. Naturk. Preuss. K. Physik.-Oekon. Ges. Koenigsberg
1: 1-102.
Habitat: Arboreal

Tribe Lasiini
Genus Euprenolepis Emery 1906
Type species: Prenolepis procera, by original designation.
Taxonomic Reference
Emery, C. 1906 Note sur Prenolepis vividula Nyl. et sur la classification des especes du genre Prenolepis. Ann.
Soc. Entomol. Belgique 50: 130-134.

Genus Paratrechina Motschoulsky 1863

Type species: Paratrechina currens, designated by Wheeler (1911).
Taxonomic Reference
Motschoulsky, V. de. 1863 Essai d'un catalogue des insectes de l'ile Ceylan. Bull. Soc. Imp. Nat. Moscou
36: 1-153.
Habitat: Nests in open soil or under rocks or other objects, or in rotten wood on the ground.
Biology: Generalized foragers

Genus Pseudolasius Emery 1887

Type species: Formica familiaris, designated by Bingham (1903).
Taxonomic Reference
Emery, C. 1887 Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza.
Formiche della regione Indo-Malese e dell'Australia. Ann. Mus. Civ. Stor. Nat. Genova (2)4(24): 209-258.
Habitat: Nest in soil under rocks or logs or in rotten wood on the ground.
Biology: Cryptic foragers. Tend homopterans

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Y. HASHIMOTO

Tribe Myrmoteratini
Genus Myrmoteras Forel 1893
Type species: Myrmoteras binghamii, by monotypy.
Taxonomic Reference
Forel, A. 1893 Note preventive sur un nouveau genre et une nouvelle espece de formicide (camponitide). Ann.
Soc. Entomol. Belgique 37: 607-608.
Moffett, M.W. 1985 Revision of the genus Myrmoteras. Bulletin of the Museum of Comparative Zoology
151: 1-53
Agosti, D. 1992 Revision of the ant genus Myrmoteras of the Malay Archipelago. Revue Suisse de Zoologie
99: 405-429
Habitat: Epigaeic
Biology: Preadators on forest floor

Tribe Oecophyllini
Genus Oecophylla Smith 1860
Type species: Formica virescens, designated by Bingham (1903).
Taxonomic Reference
Smith, F. 1860. Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the Islands of Bachian,
Kaisaa, Amboyna, Gilolo, and at Dory in New Guinea. J. Proc. Linn. Soc. London Zool. 5: 93-143.
Habitat: Nesting in arboreal, silk-woven leaf nests.
Biology: Predator, tend homopterans.

Tribe Plagiolepidini
Genus Acropyga Roger 1862
Type species: Acropyga (Atopodon) inezae Forel, designated by Wheeler (1913).
Taxonomic Reference
Roger, J. 1862. Einige neue exotische Ameisen-Gattungen und Arten. Berl. Entomol. Z. 6: 233-254.
Habitat: Hypogaeic
Biology: Tend coccids on the roots of plants

Genus Anoplolepis Santschi 1914

Type species: Formica longipes Jerdon, by monotypy.
Taxonomic Reference
Santschi, F. 1914. Formicidae. Voyage de Ch. Alluaud et. R. Jeannel en Afrique Orientale (1911-1912).
Resultats scientifiques. Hymenoptera 2: 41-148.
Habitat: Nesting in soil
Biology: Generalized foragers

Genus Lepisiota Santschi 1926

Type species: Plagiolepis rothneyi, by original designation.
Taxonomic Reference
Santschi, F. 1926 Trois notes myrmecologiques. Ann. Soc. Entomol. France 95: 13-28.
Biology: Generalized foragers

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Genus Plagiolepis Mayr 1861

Type species: Formica pygmaea, by monotypy.
Taxonomic Reference
Mayr, G. 1861. Die Europäschen Formiciden. (Ameisen.), Vienna. 80 pp.
Biology: Generalized foragers

Subfamily LEPTANILLINAE
Tribe Anomalomyrmini
Genus Anomalomyrma Taylor 1990, in Bolton 1990
Type species: Anomalomyrma taylori Bolton, by original designation.
Taxonomic Reference
Bolton, B. 1990. The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae).
Syst. Entomol. 15: 267-282.
Habitat: Nesting in soil
Biology: Cryptic predaters

Genus Protanilla Taylor 1990, in Bolton 1990

Type species: Protanilla rafflesi, by original designation.
Taxonomic Reference
Bolton, B. 1990 The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae).
Syst. Entomol. 15: 267-282.
Habitat: Nesting in soil or in rotten wood
Biology: Cryptic predators

Tribe Leptanillini
Genus Leptanilla Emery 1870
Type species: Leptanilla revelierii, by monotypy.
Taxonomic Reference
Emery, C. 1870. Studi mirmecologici. Boll. Soc. Entomol. Ital. 2: 193-201.
Baroni Urbani, C. 1977. Materiali per una revisione della sottofamiglia Leptanillinae Emery. Entomologica
Basiliensia 2: 427-488
Habitat: in soil
Biology: Cryptic mass predators (of centipedes)

Subfamily MYRMICINAE
Tribe Basicerotini
Genus Eurhopalothrix Brown & Kempf 1961
Type species: Rhopalothrix bolaui, by original designation.
Taxonomic Reference
Brown, W. L. and W. W. Kempf. 1961 The type species of the ant genus Eurhopalothrix. Psyche 67: 44.
Taylor, R.W. 1968 Notes on the Indo-Australian basicerotine ants. Australian Journal of Zoology 16: 333-348
Habitat: Nesting in leaf litter
Biology: Predators

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Tribe Cataulacini
Genus Cataulacus Smith 1853
Type species: Cataulacus taprobanae, designated by Bingham (1903).
Taxonomic Reference
Smith, F. 1853. Monograph of the genus Cryptocerus, belonging to the group Cryptoceridae - family Myrmicidae
- division Hymenoptera Heterogyna. Trans. Entomol. Soc. Lond. (2)2(1854): 213-228.
Bolton, B. 1974 A revision of the Palaeotropical arboreal ant genus Cataulacus F. Smith. Bulletin of the British
Museum (Natural History) (Entomology) 30: 1-105
Habitat: Arboreal, nesting in plant cavities

Tribe Crematogastrini
Genus Crematogaster Lund 1831
Type species: Formica scutellaris, designated by Bingham (1903).
Taxonomic Reference
Lund, P. W. 1831 Lettre sur les habitudes de quelques fourmis du Bresil, adressee a M. Audouin. Ann. Sci. Nat.
23: 113-138.
Habitat: Arboreal, nesting in hollow tree trunks and branches. Some in soil
Biology: Generalized foragers

Tribe Dacetonini
Genus Pyramica Roger 1862
Type species: Pyramica gundlachi, by monotypy.
Taxonomic Reference
Roger, J. 1862 Einige neue exotische Ameisen-Gattungen und Arten. Berl. Entomol. Z. 6: 233-254.
Bolton, B. 1999 Ant genera of the tribe Dacetonini (Hymenoptera: Formicidae). Journal of Natural History
33: 1639-1689
Habitat: Nesting in leaf litter
Biology: Predators, mainly of collembolans

Genus Strumigenys Smith 1860

Type species: Strumigenys mandibularis, by monotypy.
Taxonomic Reference
Smith, F. 1860. Descriptions of new genera and species of exotic Hymenoptera. J. Entomol. 1: 65-84.
Bolton, B. 1999 Ant genera of the tribe Dacetonini (Hymenoptera: Formicidae). Journal of Natural History
33: 1639-1689
Habitat: Nesting in leaf litter
Biology: Predators, mainly of collembolans

Tribe Formicoxenini
Genus Cardiocondyla Emery 1869
Type species: Cardiocondyla elegans, by monotypy.
Taxonomic Reference
Emery, C. 1869 Enumerazione dei Formicidi che rinvengonsi nei contorni di Napoli. Ann. Accad. Asp. Nat.
(2)2: 1-26.

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Habitat: Nesting in ground

Biology: Generalized forage on the ground surface

Genus Dilobocondyla Santschi 1910

Type species: Atopomyrmex selebensis, designated by Wheeler (1911).
Taxonomic Reference
Santschi, F. 1910 Deux nouvelles fourmis du Tonkin. Naturaliste (2)32: 283-284.
Habitat: Arboreal, nesting in twigs.

Genus Paratopula Wheeler 1919

Type species: Atopomyrmex ceylonicus, by original designation.
Taxonomic Reference
Wheeler, W. M. 1919 The ants of Borneo. Bull. Mus. Comp. Zool. Harvard Coll. 63: 43-147.
Bolton, B. 1988 A review of Paratopula Wheeler, a forgotten genus of myrmicine ants. Entomologist’s Monthly
Magazine 124: 125-143
Habitat: Arboreal (?)

Genus Rotastruma Bolton 1991

Type species: Rotastruma recava, by original designation.
Taxonomic Reference
Bolton, B. 1991. New myrmicine ant genera from the Oriental region (Hymenoptera: Formicidae). Syst. Entomol.
16: 1-13.
Habitat: Arboreal
Biology: Foraging on tree

Genus Vombisidris Bolton 1991

Type species: Vombisidris philax, by original designation.
Taxonomic Reference
Bolton, B. 1991. New myrmicine ant genera from the Oriental region (Hymenoptera: Formicidae). Syst. Entomol.
16: 1-13.
Habitat: Arboreal
Biology: Foraging on tree

Tribe Melissotarsini
Genus Rhopalomastix Forel 1900
Type species: Rhopalomastix rothneyi, by monotypy.
Taxonomic Reference
Forel, A. 1900 Un nouveau genre et une nouvelle espece de myrmicide. Ann. Soc. Entomol. Belgique 44: 24-26.
Habitat: Nesting under bark (?)
Biology: foraging on under bark, or on low vegetation

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Tribe Meranoplini
Genus Meranoplus Smith 1853
Type species: Cryptocerus bicolor, designated by Bingham (1903).
Taxonomic References
Smith, F. 1853. Monograph of the genus Cryptocerus, belonging to the group Cryptoceridae - family Myrmicidae
- division Hymenoptera Heterogyna. Trans. Entomol. Soc. Lond. (2)2(1854): 213-228.
Habitat: Nesting in the ground
Biology: Scavengers (some specialize on seeds)

Tribe Metaponini
Genus Liomyrmex Mayr 1865
Type species: Myrmica caeca.
Taxonomic References
Mayr, G.. 1865 Reise der Oesterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859, unter
den befehlen des Commodore B. von Wuellerstorf-Urbair. Zoologischer Theil. Formicidae., Vienna. 119 pp.
Ettershank, G. 1966 A generic revision of the world Myrmicinae related to Solenopsis and Pheidologeton.
Australian Journal of Zoology 14: 73-171
Rigato, F. & B. Bolton 2001 The ant genus Liomyrmex: a review (Hymenoptera Formicidae). Boll Soc. entomol.
ital., 133(3):247-256.
Habitat: Nesting under bark, or in rotten woods

Genus Metapone Forel 1911

Type species: Metapone greeni, by monotypy.
Taxonomic References
Forel, A. 1911 Sur le genre Metapone n.g. nouveau groupe des formicides et sur quelques autres formes nouvelles.
Rev. Suisse Zool. 19: 445-459.
Habitat: Nesting in hollow twings, or in rotten woods
Biology: Predators of termites

Genus Vollenhovia Mayr 1865

Type species: Vollenhovia punctatostriata, by monotypy.
Taxonomic References
Mayr, G.. 1865 Reise der Oesterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859, unter
den befehlen des Commodore B. von Wuellerstorf-Urbair. Zoologischer Theil. Formicidae. , Vienna. 119 pp.
Habitat: Many nesting under bark in logs
Biology: Predators (of termites)

Tribe Myrmecinini
Genus Myrmecina Curtis 1829
Type species: Myrmecina latreillii, by monotypy.
Taxonomic References
Curtis, J. 1829 British entomology; being illustrations and descriptions of the genera of insects found in Great
Britain and Ireland. Vol. 6:242-288. , London.
Habitat: Nesting in soil
Biology: Predators of mites

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Genus Acanthomyrmex Emery 1893

Type species: Acanthomyrmex luciolae Emery, designated by Bingham (1903).
Taxonomic References
Emery, C. 1893. Untitled. Bull. Bimens. Soc. Entomol. France 61: cclxxv-cclxxvii.
Moffett, M.W. 1986 Revision of the myrmicine genus Acanthomyrmex. Bulletin of the Museum of Comparative
Zoology 151: 55-89
Habitat: Nesting in cavities in logs, or in leaf letter
Biology: Seed harvesters (especially of ficus)

Genus Pristomyrmex Mayr 1866

Type species: Pristomyrmex pungens, by monotypy.
Taxonomic References
Mayr, G. 1866 Diagnosen neuer und wenig gekannter Formiciden. Verh. Zool.-Bot. Ges. Wien 16: 885-908.
Habitat: Nesting in rotten wood on the ground, or in soil under stones.
Biology: Generalized foragers.

Tribe Myrmicarini
Genus Myrmicaria Saunders 1842
Type species: Myrmicaria brunnea, by monotypy.
Taxonomic References
Saunders, W. W. 1842 Descriptions of two hymenopterous insects from northern India. Trans. Entomol. Soc.
Lond. 3: 57-58.
Habitat: Many Arboreal
Biology: Generalized foragers.

Tribe Phalacromyrmecini
Genus Ishakidris Bolton 1984
Type species: Ishakidris ascitaspis, by original designation.
Taxonomic References
Bolton, B. 1984 Diagnosis and relationships of the myrmicine ant genus Ishakidris gen.n. Systematic Entomology
9: 373-382
Habitat: Leaf litter
Note: The single species, I. ascitaspis, is known from Winkler bag sample collected in Sarawak.

Tribe Pheidolini
Genus Aphaenogaster Mayr 1853
Type species: Aphaenogaster sardoa Mayr, designated by Bingham (1903).
Taxonomic References
Mayr, G. 1853 Beitraege zur Kenntniss der Ameisen. Verh. Zool.-Bot. Ver. Wien 3: 101-114.
Habitat: Nesting in soil
Biology: Generalized foragers

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Genus Chimaeridris Wilson 1989

Type species: Chimaeridris boltoni, by original designation.
Taxonomic References
Wilson, E. O. 1989 Chimaeridris, a new genus of hook-mandibled myrmicine ants from tropical Asia (Hymenoptera:
Formicidae). Ins. Soc. 36: 62-69.

Genus Lophomyrmex Emery 1892

Type species: Oecodoma quadrispinosa, by monotypy.
Taxonomic References
Emery, C. 1892 Sopra alcune formiche raccolte all'Ingegnere L. Bricchetti Robecchi nel paese dei Somali. Ann.
Mus. Civ. Stor. Nat. Genova (2)12(32): 110-122.
Rigato, F. 1994. Revision of the myrmicine ant genus Lophomyrmex, with a review of its taxonomic position.
Systematic Entomology 19: 47-60
Habitat: Nesting in soil near the base of trees
Biology: General foragers

Genus Pheidole Westwood 1839

Type species: Atta providens, by monotypy.
Taxonomic References
Westwood, J. O. 1839. An introduction to the modern classification of insects Vol. 2, part 11, pp. 193-224. , London.
Eguchi, K. 2001. A revision of the Bornean species of the ant genus Pheidole (Insects: Hymenoptera: Formicidae:
Myrmicinae). Tropics Monograph Series 2: 1-154
Habitat: Nesting in soil, or in rotten wood
Biology: General foragers

Tribe Pheidologetonini
Genus Oligomyrmex Mayr 1867
Type species: Oligomyrmex concinnus, by monotypy.
Taxonomic References
Mayr, G. 1867. Adnotationes in monographiam formicidarum Indo-Neerlandicarum. Tijdschr. Entomol. (2)2(10):
33-117.
Habitat: in soil
Biology: Prey on brood and eggs of termites, or ants

Genus Pheidologeton Mayr 1862

Type species: Oecodoma diversa, designated by Bingham (1903).
Taxonomic References
Mayr, G. 1862 Myrmecologische Studien. Verh. Zool.-Bot. Ges. Wien 12: 649-776.
Habitat: Nesting in soil
Biology: Generalized and mass foragers

Genus Recurvidris Bolton1890

Type species: Trigonogaster recurvispinosa Forel, by monotypy. Replacement name for Trigonogaster Forel, by
Bolton (1992).

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Taxonomic References
Forel, A. 1890. Aenictus-Typhlatta decouverte de M. Wroughton. Nouveaux genres de formicides. Ann. Soc.
Entomol. Belg. Comptes-rendus Seances 34: cii-cxiv.
Bolton, B. 1992. A review of the ant genus Recurvidris, a new name for Trigonogaster Forel. Psyche 99: 35-48
Habitat: Nesting in soil
Biology: Generalized foragers

Tribe Solenopsidini
Genus Anillomyrma Emery 1913
Type species: Monomorium decamerum Emery, by monotypy.
Taxonomic References
Emery, C. 1913 Etudes sur les Myrmicinae. Ann. Soc. Entomol. Belgique 57: 250-262.
Bolton, B. 1987 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr
(Hymenoptera: Formicidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 54: 263-452.
Habitat: Nesting in soil, or in termite nests
Biology: Prey on termites (?)

Genus Epelysidris Bolton 1987

Type species: Epelysidris brocha, by original designation.
Taxonomic References
Bolton, B. 1987. A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr
(Hymenoptera: Formicidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 54: 263-452.
Habitat: Moss forest, nesting in soil under the moss and rocks.
Note: Only single species, E. brocha, is known form Sarawak.

Genus Monomorium Mayr 1855

Type species: Monomorium minutum, by monotypy.
Taxonomic References
Mayr, G. 1855 Formicina austriaca. Beschreibung der bisher im oesterreichischen Kaiserstaate aufgefundenen
Ameisen nebst Hinzufuegung jener in Deutschland, in der Schweiz und in Italien vorkommenden Ameisen. Verh.
Zool.-Bot. Ver. Wien 5: 273-478.
Bolton, B. 1987 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr
(Hymenoptera: Formicidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 54: 263-452.
Habitat: Nesting under rocks, under bark, or in rotten branches.
Biology: Scavengers. Seed harvesters

Genus Solenopsis Westwood 1840

Type species: Solenopsis mandibularis, by monotypy.
Taxonomic References.
Westwood, J. O. 1840 Observations on the genus Typhlopone, with descriptions of several exotic species of ants.
Ann. Mag. Nat. Hist. 6: 81-89.
Bolton, B. 1987 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr
(Hymenoptera: Formicidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 54: 263-452.
Habitat: Nesting in ground
Biology: Generalized foragers and thief ants

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Y. HASHIMOTO

Tribe Stenammini
Genus Calyptomyrmex Emery 1887
Type species: Calyptomyrmex beccarii Emery, by monotypy.
Taxonomic References
Emery, C. 1887 Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza.
Formiche della regione Indo-Malese e dell'Australia (continuazione e fine). Ann. Mus. Civ. Stor. Nat. Genova
(2)5(25): 427-473.
Baroni Urbani, C. 1975 Primi reperti del genere Calyptomyrmex Emery nel subcontinente Indiano. Entomologica
Basiliensia 1: 395-411
Habitat: Leaf litter

Genus Dacetinops Brown & Wilson 1957

Type species: Dacetinops cibdelus, by original designation.
Taxonomic References
Brown, W. L. and E. O. Wilson. 1957. Dacetinops, a new ant genus from New Guinea. Breviora 77: 1-7.
Taylor, R.W. 1965. A new species of the ant genus Dacetinops from Sarawak. Breviora 237: 1-4.
Taylor, R.W. 1985. The ants of the Papuasian genus Dacetinops. In Ball, G.E. (Ed.). Taxonomy, Phylogeny and
Zoogeography of Beetles and Ants. 41-67

Genus Lasiomyrma Terayama & Yamane 2000

Type species: Lasiomyrma gedensis Terayama & Yamane, by original designation
Taxonomic References
Terayama, M. and S. Yamane 2000. Lasiomyrma, a new stenammine ant genus from Southeast Asia (Hymenoptera:
Formicidae) Entomological Science 3(3): 523-527.
Habitat: Nesting in rotten woods

Genus Lordomyrma Emery 1897

Type species: Lordomyrma furcifera, designated by Wheeler (1911).
Taxonomic References
Emery, C. 1897 Formicidarum species novae vel minus cognitae in collectione Musaei Nationalis Hungarici, quas
in Nova Guinea, colonia germanica, collegit L. Biro. Term. Fuz. 20: 571-599.
Habitat: Nesting in soil, or in rotten woods

Genus Mayriella Forel 1902

Type species: Mayriella abstinens, by monotypy.
Taxonomic References
Forel, A. 1902. Fourmis nouvelles d'Australie. Rev. Suisse Zool. 10: 405-548.
Habitat: Nesting in soil, or rotten logs
Biology: Foraging in leaf litter

Genus Proatta Forel 1912

Type species: Proatta butteli, by monotypy.
Taxonomic References
Forel, A. 1912 Descriptions provisoires de genres, sous-genres et especes de formicides des Indes orientales. Rev.

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Suisse Zool. 20: 761-774.

Habitat: Nesting in soil
Biology: Scavengers

Genus Tetheamyrma Bolton 1991

Type species: Tetheamyrma subspongia, by original designation.
Taxonomic References
Bolton, B. 1991 New myrmicine ant genera from the Oriental region (Hymenoptera: Formicidae). Syst. Entomol.
16: 1-13.
Habitat: Nesting in leaf letter
Note: Only single species, T. subspongia, is known from Sabah, Poring.

Tribe Tetramoriini
Genus Rhoptromyrmex Mayr 1901
Type species: Rhopalomastix rothneyi, by monotypy.
Taxonomic References
Mayr, G. 1901. Suedafrikanische Formiciden, gesammelt von Dr. Hans Brauns. Ann. Naturhistor. Mus. Wien
16: 1-30.
Bolton, B. 1986 A taxonomic and biological review of the tetramoriine ant genus Rhoptromyrmex. Systematic
Entomology 11: 1-17
Habitat: Nesting in soil
Biology: Social parasites (of Pheidole, Tapinoma)

Genus Secostruma Bolton 1988

Type species: Secostruma lethifera, by original designation.
Taxonomic References
Bolton, B. 1988. Secostruma, a new subterranean tetramoriine ant genus (Hymenoptera: Formicidae). Syst. Entomol.
13: 263-270.
Habitat: Nesting in soil
Note: Only single species, S. lethifera, is known from soil core sample collected in Sabah, Gunong Silam.

Genus Tetramorium Mayr 1855

Type species: Formica caespitum, designated by Girard (1879).
Taxonomic References
Mayr, G. 1855. Formicina austriaca. Beschreibung der bisher im oesterreichischen Kaiserstaate aufgefundenen
Ameisen nebst Hinzufuegung jener in Deutschland, in der Schweiz und in Italien vorkommenden Ameisen. Verh.
Zool.-Bot. Ver. Wien 5: 273-478.
Bolton, B. 1977. The ant tribe Tetramoriini. The genus Tetramorium Mayr in the
Oriental and Indo-Australian regions, and in Australia. Bulletin of the British Museum (Natural History)
(Entomology) 36: 67-151
Habitat: Nesting in soil
Biology: Generalized foragers

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Y. HASHIMOTO

Subfamily PONERINAE
Tribe Amblyoponini
Genus Amblyopone Erichson 1842
Type species: Amblyopone australis Erichson, by monotypy.
Taxonomic References
Erichson, W. F. 1842. Beitrag zur Insecten-Fauna von Vandiemensland, mit besonderer Beruecksichtigung der
geographischen Verbreitung der Insecten. Arch. Naturgesch. 8: 83-287.
Brown, W.L.,Jr. 1960. Contributions toward a reclassification of the Formicidae. 3. Tribe Amblyoponini. Bulletin
of the Museum of Comparative Zoology at Harvard College 122: 145-230
Habitat: nesting in soil under rocks or logs, or in rotten wood
Biology: Cryptic predators (some specialist to prey centipeds)

Genus Myopopone Roger 1861

Type species: Myopopone maculata, designated by Bingham (1903).
Taxonomic References
Roger, J. 1861. Die Ponera-artigen Ameisen. (Schluss.) Berl. Entomol. Z. 5: 1-54.
Brown, W.L.,Jr. 1960 Contributions toward a reclassification of the Formicidae. 3. Tribe Amblyoponini. Bulletin
of the Museum of Comparative Zoology at Harvard College 122:145-230
Habitat: Nesting in rotten wood or under bark.
Biology: Predator (some specialist predator on millipeds)

Genus Mystrium Roger 1862

Type species: Mystrium mysticum, by monotypy.
Taxonomic References
Roger, J. 1862. Einige neue exotische Ameisen-Gattungen und Arten. Berl. Entomol. Z. 6: 233-254.
Brown, W.L.,Jr. 1960 Contributions toward a reclassification of the Formicidae. 3. Tribe Amblyoponini. Bulletin
of the Museum of Comparative Zoology at Harvard College 122: 145-230
Habitat: Nesting under rocks or logs on the ground and in leaf litter.
Biology: Predator

Genus Prionopelta Mayr 1866

Type species: Prionopelta punctulata, by monotypy.
Taxonomic References
Mayr, G. 1866. Myrmecologische Beitraege. Sitzungsber. K. Wiss. Math.-Naturwiss. Classe 53: 484-517.
Brown, W.L.,Jr. 1960 Contributions toward a reclassification of the Formicidae. 3. Tribe Amblyoponini. Bulletin
of the Museum of Comparative Zoology at Harvard College 122: 145-230
Habitat: Nesting under rocks or logs on the ground and in leaf litter.
Biology: Predator (of chilopda)

Tribe Ectatommini
Genus Discothyrea Roger 1863
Type species: Discothyrea testacea, by monotypy.
Taxonomic References
Roger, J. 1863 Die neu aufgefürten Gattungen und Arten meines Formiciden-Verzeichnisses, nebst Ergäzung

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einiger früer gegeben Beschreibungen. Berl. Entomol. Z. 7: 131-214.

Brown, W.L.,Jr. 1958 Contributions toward a reclassification of the Formicidae. 2. Tribe Ectatommini. Bulletin
of the Museum of Comparative Zoology at Harvard College 118: 175-362
Habitat: Nesting in open soil, or in soil under rocks, or in rotten wood.
Biology: Specialist predators on arthropod eggs.

Genus Gnamptogenys Roger 1863

Type species: Ponera tornata, designated by Emery (1911).
Taxonomic References
Roger, J. 1863 Die neu aufgefürten Gattungen und Arten meines Formiciden-Verzeichnisses, nebst Ergäzung
einiger früer gegeben Beschreibungen. Berl. Entomol. Z. 7: 131-214.
Brown, W.L.,Jr. 1958 Contributions toward a reclassification of the Formicidae. 2. Tribe Ectatommini. Bulletin
of the Museum of Comparative Zoology at Harvard College 118: 175-362
Habitat: Nesting in rotten logs on the ground.
Biology: Predators (some specialist predators on millipeds)

Genus Proceratium Roger 1863

Type species: Proceratium silaceum, by monotypy.
Taxonomic References
Roger, J. 1863 Die neu aufgef¸rten Gattungen und Arten meines Formiciden-Verzeichnisses, nebst Erg‰zung
einiger fr¸er gegeben Beschreibungen. Berl. Entomol. Z. 7: 131-214.
Brown, W.L.,Jr. 1958 Contributions toward a reclassification of the Formicidae. 2. Tribe Ectatommini. Bulletin
of the Museum of Comparative Zoology at Harvard College 118: 175-362
Habitat: Nesting in soil or rotten wood.
Biology: Specialist predators of arthropod eggs.

Tribe Platythyreini
Genus Platythyrea Roger 1863
Type species: Pachycondyla punctata, designated by Bingham (1903).
Taxonomic References
Roger, J. 1863 Die neu aufgefürten Gattungen und Arten meines Formiciden-Verzeichnisses, nebst Ergäzung
einiger früer gegeben Beschreibungen. Berl. Entomol. Z. 7: 131-214.
Brown, W. L., Jr. 1975 Contributions toward a reclassification of the Formicidae. V. Ponerinae, tribes Platythyreini,
Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini. Search Agric. (Ithaca N. Y.) 5(1): 1-115
Habitat: Nesting in soil, in rotten wood or in hollow twigs on trees
Biology: Predator (some specialist predators on termites)

Genus Probolomyrmex Mayr 1901

Type species: Probolomyrmex filiformis, by monotypy.
Taxonomic References
Mayr, G. 1901 Suedafrikanische Formiciden, gesammelt von Dr. Hans Brauns. Ann. Naturhistor. Mus. Wien
16: 1-30.
Taylor, R.W. 1965 A monographic revision of the rare tropicopolitan ant genus Probolomyrmex Mayr. Transactions
of the Royal Entomological Society of London 117: 345-365

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Y. HASHIMOTO

Brown, W. L., Jr. 1975 Contributions toward a reclassification of the Formicidae. V. Ponerinae, tribes Platythyreini,
Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini. Search Agric. (Ithaca N. Y.) 5(1): 1-115
Habitat: Nesting in soil under rocks, in leaf litter or in rotten wood.
Biology: Cryptic predator

Tribe Ponerini
Genus Anochetus Mayr 1861
Type species: Odontomachus ghilianii Spinola, by monotypy.
Taxonomic References
Mayr, G.. 1861 Die Europ¸schen Formiciden. (Ameisen.) , Vienna. 80 pp.
Brown, W.L.,Jr. 1978 Contributions toward a reclassification of the Formicidae. Part 6. Ponerinae, tribe Ponerini,
subtribe Odontomachiti. Section B. Genus Anochetus and bibliography. Studia Entomologica (N.S.) 20: 549-652
Habitat: Nesting in soil, in termite nests or under logs.
Biology: Predator

Genus Centromyrmex Mayr 1866

Type species: Centromyrmex bohemanni, by monotypy
Taxonomic References
Mayr, G. 1866 Diagnosen neuer und wenig gekannter Formiciden. Verh. Zool.-Bot. Ges. Wien 16: 885-908.
Habitat: Nesting in soil (in or near termite nest)
Biology: Predator of termite

Genus Cryptopone Emery 1893

Type species: Cryptopone testacea, by monotypy.
Taxonomic References
Emery, C. 1893 Untitled. Bull. Bimens. Soc. Entomol. France 61: cclxxv-cclxxvii.
Brown, W.L.,Jr. 1963. Characters and synonymies among the genera of ants. Part 3. Some members of the tribe
Ponerini. Breviora 190: 1-10
Habitat: Nesting in soil under rocks, or in rotten wood.
Biology: Cryptic predator

Genus Diacamma Mayr 1862

Type species: Ponera rugosa, designated by Bingham (1903).
Taxonomic References
Mayr, G. 1862 Myrmecologische Studien. Verh. Zool.-Bot. Ges. Wien 12: 649-776.
Habitat: Nesting in soil
Biology: Predator

Genus Emeryopone Forel 1912

Type species: Emeryopone buttelreepeni, by monotypy.
Taxonomic References
Forel, A. 1912 Descriptions provisoires de genres, sous-genres et especes de formicides des Indes orientales.
Rev. Suisse Zool. 20: 761-774.
Baroni Urbani, C. 1975 Contributo alla conoscenza dei generi Belonopelta Mayr e Leiopelta gen.n. Mitteilungen

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der Schweizerischen Entomologischen Gesellschaft 48: 295-310

Habitat: Nesting in soil
Biology: Predator

Genus Harpegnathos Jerdon 1851

Type species: Harpegnathos saltator, by monotypy.
Taxonomic References
Jerdon, T. C. 1851 A catalogue of the species of ants found in southern India. Madras J. Lit. Sci. 17: 103-127.
Habitat: Nesting in soil
Biology: Predator

Genus Hypoponera Santschi 1938

Type species: Ponera abeillei, by original designation.
Taxonomic References
Santschi, F. 1938 Notes sur quelques Ponera. Bull. Soc. Entomol. France 43: 78-80.
Habitat: Nesting in soil, in rotten wood, in leaf litter.
Biology: Cryptic predators (some specialist predator on Collembola).

Genus Leptogenys Roger 1861

Type species: Leptogenys falcigera, designated by Bingham (1903).
Taxonomic References
Roger, J. 1861 Die Ponera-artigen Ameisen. (Schluss.) Berl. Entomol. Z. 5: 1-54.
Habitat: Nesting in soil, in rotten wood, in leaf litter.
Biology: Mass foraging predator (some having army ant like habit)

Genus Myopias Roger 1861

Type species: Myopias amblyops, by monotypy.
Taxonomic References
Roger, J. 1861 Die Ponera-artigen Ameisen. (Schluss.) Berl. Entomol. Z. 5: 1-54
Habitat: Nesting in soil, or in rotten wood
Biology: Predator

Genus Odontomachus Latreille 1804

Type species: Formica haematoda, by monotypy.
Taxonomic References
Latreille, P. A. 1804 Tableau methodique des insectes. Classe huitieme. Insectes, Insecta. Nouveau Dictionnaire
d'Histoire Naturelle 24: 129-200.
Brown, W.L., Jr. 1976 Contributions toward a reclassification of the Formicidae. Part 6. Ponerinae, tribe Ponerini,
subtribe Odontomachiti. Section A. Introduction, subtribal characters, genus Odontomachus.Studia Entomologica
(N.S.) 19: 67-171
Habitat: Nesting in soil, or in rotten wood on the ground.
Biology: Predator

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 Y. HASHIMOTO

Genus Odontoponera Mayr 1862

Type species: Ponera denticulata, by monotypy.
Taxonomic References
Mayr, G. 1862 Myrmecologische Studien. Verh. Zool.-Bot. Ges. Wien 12: 649-776.
Habitat: Nesting in soil
Biology: Predator

Genus Pachycondyla Smith 1858

Type species: Formica crassinoda, designated by Emery (1901).
Taxonomic References
Smith, F.. 1858 Catalogue of the hymenopterous insects in the collection of the British Museum. 6. Formicidae. ,
London. 216 pp.
Habitat: Nesting in soil, or in logs
Biology: Predator, scavengers

Genus Ponera Latreille 1804

Type species: Formica coarctata, designated by Westwood (1840).
Taxonomic References
Latreille, P. A. 1804 Tableau methodique des insectes. Classe huitieme. Insectes, Insecta. Nouveau Dictionnaire
d'Histoire Naturelle 24: 129-200.
Taylor, R.W. 1967 A monographic revision of the ant genus Ponera Latreille. Pacific Insects Monograph
13: 1-112
Habitat: Nesting in the soil, in rotten wood, or under bark or moss on rotten logs.
Biology: Cryptic predators

Subfamily PSEUDOMYRMECINAE
Tribe Pseudomyrmecini
Genus Tetraponera Smith 1852
Type species: Tetraponera atrata, designated by Wheeler (1911).
Taxonomic References
Smith, F. 1852 Descriptions of some hymenopterous insects captured in India, with notes on their oeconomy, by
Ezra T. Downes, Esq., who presented them to the Honourable the East India Company. Ann. Mag. Nat. Hist.
(2)9: 44-50.
Ward, P. S. 2001 Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae)
in the Oriental and Australian regions. Invert. Taxon. 15: 589-665.
Habitat: Arboreal, nesting in plants cavities
Biology: Predator (?)

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