Philippine Ants

This page is intended to promote international collaboration on the study of the ants of the Philippines. David M.General is working in the Philippines and is focusing on the biodiversity of Philippine ants. Gary D. Alpert is at Harvard University, Museum of Comparative Zoology and is interested in the biodiversity of ants from the Philippines. Perry Buenavente, Museum Researcher from the Entomology Section, Zoology Division, National Museum of Natural History, Philippine National Museum is working on the biodiversity of Philippine ants. Robert Taylor, adjunct professor at the Australian National University is working on the taxonomy of ant genera many of which are found in the Philippines. Others interested in this topic are welcome to join this AntWiki group.

The richness of the Philippine ant fauna has not been fully examined. There are many areas that have not been inventoried for ants and surveying these areas will yield many new species records.We currently know that 489 valid species and subspecies occur in the Philippines. This species list is based upon the effort of many ant collectors as well as myrmecologists who have published on the taxonomy of Philippine ants. Those who have specimen records that expand this list are encouraged to submit this information in the discussion section of the Philippine Ants web page.

Lists of Taxa

List of Philippine Ant Species, with images for each species.

The Ants of the Philippines

This project includes all genera and species of ants found in the Philippines.

Abstract

An overview of the history of myrmecology in the Philippine archipelago is presented. Keys are provided to the 11 ant subfamilies and to the 92 ant genera known from the Philippines. Thirteen ant genera (14%), including 3 undescribed genera, are recorded for the first time from the Philippines. The biology and ecology of the 92 ant genera of the Philippines are summarized in the form of brief generic accounts. A bibliography of significant taxonomic and behavioral papers on Philippine ants and a checklist of valid species and their island distribution are provided.

Introduction

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  Montane Forest, Mt. Palali, Luzon Island.

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  Slash and burn agriculture, Luzon Island.

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  Beach Forest, Aurora Province, Luson Island.

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  Deforested mountain, northwestern Mindoro Island.

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  Mossy Forest, Mt. Balatukan, Mindanao Island.

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  Lowland Dipterocarp Forest, Bucas Grande Island, Surigao del Norte Province.

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  Lowland Dipterocarp Forest, Puerto Princesa City, Palawan.

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  Limestone Karst Forest, Puerto Princesa City, Palawan.

The study of ants can be difficult, particularly in the tropics. Tropical ant faunas are only partly explored and information concerning them is widely scattered throughout the scientific literature. Taxonomic references are frequently hard to access, badly outdated, and difficult for the non-specialist to use. The systematics of many of the major genera (e.g., Camponotus, Crematogaster, Pachycondyla) is still chaotic and unrecognized synonyms and undescribed species abound. Taxonomic progress is relatively slow because most type material of described material is deposited in Europe, North America, or Japan, and museums are understandably hesitant to send fragile types to workers in the developing world (Naskrecki 2004). Introductory accounts of important tropical ant faunas are badly needed to provide an entry point for biologists interested in studying ants. The purpose of this paper is to provide such an introduction to the ant fauna of the Philippines. Simplified keys to subfamilies and to genera are presented. Most character states in the keys are discernible with a 40X stereomicroscope. Technical terms and obscure character states are minimized. Full-face and profile color photo-images are included to illustrate the different genera found in the Philippines. A glossary is also included to help the student understand the precise technical terms used in the keys. And the most up-to-date systematic arrangement of genera is used (Bolton et al. 2006, Bolton 2010). We hope that this paper will stimulate local interest in the study of Philippine ants. There is much to be done and discovered, and many opportunities await the Filipino myrmecologist. The Philippines is so ecologically diverse that it is difficult to characterize the country in a single paragraph. While the climate is generally tropical and maritime, there are zones which are distinctly different. The mean temperature ranges from 25.5° C in January to 28.3° C in May, however, high-altitude locations (>1,000 masl) are sub-temperate. Baguio City, in northern Luzon, has a mean annual temperature of only 18.3˚C (PAGASA 2010). Some parts, such as southern Cotabato Province, Mindanao Island, are relatively dry with an average annual rainfall of only 978 mm (PAGASA 2010). In contrast, 5713 mm of rainfall was measured at 1650 masl on Mt. Isarog in 114 days from November 1993 to May 1994 (Heaney et al. 1999). There are also wet tropical zones, mainly in the eastern part of the country, and dry tropical zones that are in the rain shadow (west) of mountain ranges. Camarines Sur, a province in the Bicol Peninsula of Luzon Island, for instance, has three distinct climate patterns, depending on altitude and position relative to Mt. Isarog, the dominant feature of the landscape. There is also a typhoon corridor, mainly in the eastern part of the archipelago, which is visited by an average of 19 typhoons every year (PAGASA 2010). The Philippines is composed of over 7,100 islands, most of which are uninhabited. There are islands, such as Sibuyan Island in the central Philippines, which have never been connected to larger islands, even during the last Ice Age (Heaney et al. 1999). Single mountains, usually volcanoes, also exist as island habitats within the broader lowlands. This complex blend of climatic diversity and opportunity for geographic isolation has likely led to high levels of endemism, a phenomenon that deserves much further exploration. For example, two adjacent islands, Biliran and Samar, each have their own species of Meranoplus, a ground-dwelling myrmicine ant (Schödl 1998; DMG, unpublished notes). The Palawan island group is unique in that many of its plants and animals are found nowhere else in the Philippines, or in the world, for that matter. Perhaps the best documented ant fauna in the Philippines is that of the Cuernos de Negros Mountains near Dumaguete City. This area was the favorite hunting ground of the late Dr. James W. Chapman, who collected there for about two decades (Chapman and Chapman 1947). Nevertheless, it is almost certain that more intensive and systematic collection methods will turn up new species there. There are many interesting study sites in the Philippines. Particularly interesting are: old growth forests, old mangrove areas, small islands (preferably uninhabited), forest canopies, wooded ravines, reforested areas, and even microhabitats such as soil and forest canopies. Each island or province has its own opportunities. Surprisingly, new species have been found in even the most disturbed localities, such as university campuses (DMG, unpublished notes), possibly relicts of a more diverse ant assemblage (S. Cover, pers. comm.).

A short history of Philippine myrmecology

The study of Philippine ant fauna began in the late 1800’s, during the last century of the Spanish colonial period. Small numbers of Philippine ants were collected by European travelers who then sold or gave their collections to the great myrmecologists of that era, notably Auguste Forel and Carlo Emery. Emery described ants collected in Manila and Antipolo, Luzon Island by E. Simon, a Frenchman (Emery 1893c). Frederick Smith described a few species from the Philippines without specifying the provenance of the specimens (F. Smith 1858). These researchers described some new species, but made no effort to characterize the Philippine ant fauna as a whole. During the early American colonial era, Richard.C. MacGregor, a US-trained Australian ornithologist, Charles S. Banks and other biologists working for the Bureau of Science, Manila (Van Steenis-Kruseman 2006), traveled the archipelago and gave or sold their insect collections to Americans like Dr. W.H. Ashmead of the United States National Museum (USNM) and the great Dr. William Morton Wheeler of Harvard University. Ashmead described new species collected by Dr. P.L. Stangl of the U.S. Army, Prof. L.E. Griffin of Missouri Valley University, Dr. E.B. Copeland of the Government Laboratories in Manila, and Dr. M.H. Smith of the US Fish Commission (Ashmead 1904a, 1904b). Dr. Francis X. Williams, who studied under W. M. Wheeler, collected ants while he was looking for insect predators of agricultural pests, as a researcher for the USDA. He took advantage of the expertise of Dr. Charles F. Baker, a professor at the University of the Philippines College of Agriculture at Los Baños, Laguna. Baker collected insects prodigiously, amassing hundreds of Schmitt boxes, which were bequeathed to USNM (Evans 1985). The first intensive ant collecting was done by Dr. James W. Chapman, Wheeler’s colleague in Harvard, who arrived in the 1910's to be a missionary and teacher at Silliman University in Dumaguete, Negros Oriental, Negros Island. He concentrated on studying the ants of the Cuernos de Negros Mountains, but also collected in northern Luzon and Mindanao. During the Second World War, Dr. Chapman continued to collect ants even as Japanese soldiers were hunting him down. Fortunately, he survived capture and incarceration, and his collection, which he had hidden, was largely intact (Chapman and Chapman 1947). Most of these specimens are deposited in the Museum of Comparative Zoology (MCZ) at Harvard University. Chapman later published a list of the ants of Asia (Chapman and Capco 1951). Interestingly, Jesuit priests were important early collectors of Hymenoptera, including ants. Ashmead studied the specimens collected by Fr. W.A. Stanton, S.J. and Fr. Robert E. Brown, S.J. (Ashmead 1904a, 1904b). Fr. Brown collected the holotype queen of the controversial genus Pseudaphomomyrmex, recently synonymized under the genus Tapinoma (Ashmead 1905, LaPolla and Longino 2006, Fisher and Bolton 2007). Fr. B.B. Lowery S.J. also collected ants in the Philippines in the 1960s (Ward 2001, Shattuck, pers. comm.). His collections have been studied by researchers all over the world. During this period and after the war, some Filipinos also collected ants. This fact can be gleaned from the locality labels on specimens in certain museums. Specimens collected by Domingo Empeso, H. M. Torrevillas, A. Reyes, and M. Ramos are in the collections of the MCZ and the Bernice P. Bishop Museum in Hawaii. In 1966, Dr.Clare Baltazar published a monograph on Philippine Hymenoptera, which included 235 entries for ant species (Baltazar 1966). Much has changed since then. Many of the genera have been revised, many species have been added to the list, and a number of names have been synonymized. This paper presents an updated list containing 428 valid names for ants in the Philippines. There has been some recent taxonomic progress. Recent monographs relevant to the Philippine ant fauna include: Acanthomyrmex (Moffett 1986, Agosti 1992), Acropyga (LaPolla 2004), Anillomyrma (Eguchi et al. 2010), Calyptomyrmex (Shattuck 2011), Cardiocondyla (Seifert 2003), Carebara (Fernandez 2004), Euprenolepis (LaPolla 2009), Forelophilus (Zettel and Zimmerman 2007), Gnamptogenys of the Old World (Lattke 2004), Iridomyrmex (Shattuck 1992a), Leptomyrmex (Lucky and Ward 2010), Liomyrmex (Rigato and Bolton 2001), Lophomyrmex (Rigato 1994), Mayriella (Shattuck and Barnett 2007), Meranoplus (Schödl 1998), Monomorium (Heterick 2001), Myrmoteras (Moffett 1985, Agosti 1992, Zettel and Sorger 2011), Mystrium (Bihn and Verhaagh 2007), Odontomachus (Sorger and Zettel 2011), Paratopula (Bolton 1988), Prionopelta (Shattuck 2008a), Pristomyrmex (Wang 2003, Zettel 2006), Proceratium (Baroni Urbani and de Andrade 2003), Probolomyrmex (Eguchi et al. 2006), Pyramica (Bolton 2000), Recurvidris (Bolton 1992, Zettel 2008), Rhoptromyrmex (Bolton 1986), Strumigenys (Bolton 2000), Technomyrmex (Bolton 2007), Tetraponera (Ward 2001) and Vombisidris (Zettel and Sorger 2010a). Unfortunately, there has been relatively little intensive collecting, or the use of ants in ecological research, since the efforts of Chapman. A recent transect study (Alpert and General in preparation) surveyed the ants of a 27-year-old narra (Pterocarpus indicus Willd., Fabaceae) reforestation project in the Bicol region of Luzon Island. New generic records and new species are recorded. Inventories for undergraduate and graduate theses remain an important source of specimens, especially in new study sites. Fortunately, the collection methods are now more standardized (Agosti et al. eds. 2000). All specimens collected on Philippine soil and exported should have the required permits from the Philipine government.

Synopsis of Philippine ant genera

In this paper, the acronym “PH” is used to represent the archipelago of the Republic of the Philippines, in compliance with the International Organization for Standardization 3166-1 country codes (ISO, 2010). The acronym “PI” is an anachronism and should no longer be used since it refers to the islands before the country gained independence in 1946. The first electronic checklist of Philippine ants is available online at Discoverlife (Alpert et al. 2006). Another online resource is the Philippine page on AntWeb, hosted by the California Academy of Sciences (Alpert and General 2008).

Figure 1. Full-face and profile images of Philippine ant genera. Acanthomyrmex mindanao Moffett, 1986, major worker (A, B). Acropyga pallida (Donisthorpe, 1938), minor worker (C, D). Aenictus ceylonicus (Mayr, 1866) (E, F).

Acanthomyrmex

Genus Acanthomyrmex Emery, 1893a –Myrmicinae: 17 spp, 1 known from PH.

The single known valid Philippine species, Acanthomyrmex mindanao was described from specimens collected under bark in Mt. McKinley, Davao Region, southern Mindanao Island. Other specimens were collected in northern Mindanao Island: Momungan, Olangon, and Iligan City, Lanao del Sur Province and Gingoog City, Misamis Oriental Province. This species is also known from Borneo and Sarawak, Malaysia. These ants may be collected by sifting leaf litter, baiting with small seeds, and looking under bark. Keys to species: Moffett (1986) (World, revision), Terayama et al. 1998.

Acropyga

Genus Acropyga Roger, 1862 – Formicinae: 40 spp., 3 known from PH.

(New record). This genus is pantropical. These tiny, yellow, hypogaeic ants are known to tend subterranean coccids. One species known from the Philippines, Acropyga pallida, was originally described from New Guinea. These ants may be collected by sifting leaf litter and soil cores up to about 10 cm deep. Key to species: LaPolla (2004).

Aenictus

Genus Aenictus Shuckard, 1840 – Aenictinae: 149 spp., 15 known from PH.

This genus is widespread throughout the archipelago. Large colonies forage for insect prey in the leaf litter. These are true army ants, making bivouacs in tree hollows and other protected cavities. These ants may be collected by searching for conspicuous columns of emigrating or foraging ants, and carefully inspecting cavities in trees and logs. For raiding behavior, see Schneirla and Reyes (1966). Keys to species: Wheeler (1930c) (Philippines), Wilson (1964) (Indo-Australian, revision), Terayama and Yamane (1989) (Indonesia, Sumatra), Shattuck (2008b) (Australian, review of genus), Zettel and Sorger (2010b) (new Philippine species described).

Anillomyrma

Genus Anillomyrma Emery, 1913b – Myrmicinae: 2 spp., 1 known from PH.

Specimens of Anillomyrma decamera (Emery, 1901) were recently collected by Joanaviva Caceres-Plopenio in a transect study on Mt. Isarog, Luzon Island (Eguchi et al. 2010). These pale, tiny, thin-skinned subterranean ants seem to prefer sandy soils. These ants may be collected by sifting soil cores, preferably by Berlese or Winkler extraction, or baiting underground in sandy loam areas. Key to species: Eguchi et al. 2010 (World revision).

Anochetus

Genus Anochetus Mayr, 1861 – Ponerinae: 87 spp., 7 known from PH.

These small, fast-moving trap-jawed ants are general predators and usually found foraging singly on the ground or in the leaf litter. These ants are similar to Odontomachus, except that they are smaller and have blunt petiolar nodes. Key to species: Brown (1978) (World).

Key to the Anochetus of the Philippines

Zettel (2102):

At least eleven species of Anochetus have been recorded from the Philippines. One additional unnamed species of the A. longifossatus species group may belong to Anochetus schoedli. Of the eleven species, Anochetus graeffei and Anochetus ruginotus belong to the A. graeffei group, Anochetus isolatus to the A. cato group, Anochetus schoedli to the A. longifossatus group, and Anochetus princeps to the A. rugosus group. The other six species belong to the A. risii group, which shows a great diversity in southeastern Asia and eastwards to Taiwan, the Philippines and New Guinea (Brown 1978, Terayama 1989). Endemism is chiefly found in the A. risii group: Five species are only recorded from single locations in the Philippines, the identification of the sixth species (Anochetus modicus from Negros) needs confirmation. The mountainous species A. schoedli is probably also a locally endemic species, as are all other members of the A. longifossatus species group (see Brown 1978). Their small body size, light pigmentation, and minute eyes indicate a life in the lower leaf litter and possibly low dispersal abilities. Three species, A. graeffei, A. princeps and A. isolatus have wide distribution. However, more detailed taxonomic studies may change the concept of species with high morphological variability like A. graeffei and A. isolatus. Anochetus ruginotus is presently only recorded from the Philippines, but as the taxon was hitherto in synonymy with A. graeffei, it may have a wider distribution than presently known.

Considering the fragmentary collection data from a very few, well-studied sites, it can be expected that the Philippine Anochetus fauna is much richer than presently known. Preliminary results indicate a high rate of endemic species, some being possibly restricted to individual islands or mountain ranges. The situation resembles that of Odontomachus and Pristomyrmex (see Zettel 2006, Sorger and Zettel 2011).

Habitats: Most Philippine records of Anochetus are from forested slopes of mountains at low elevations. Exceptional is one lowland record of A. graeffei from Masbate, which fits well to observations on some other widely distributed, euryoecious ants species (e.g., Odontomachus simillimus; see Sorger and Zettel 2011). Wilson (1959: p. 507) speculates that “very probably it [A. graeffei] has been distributed through part of this range by the inadvertent agency of man.” Highest Anochetus records in the Philippines are the type localities of A. werneri (ca. 1000 m a.s.l.), A. schoedli (ca. 1400 - 1500 m a.s.l.) and A. brevis (1520 - 1830 m a.s.l.). Brown (1978) reports a collection of A. princeps at an elevation of ca. 1500 m (5000 ft) on Borneo. These are among the highest records in the genus.

Anoplolepis

Genus Anoplolepis Santschi, 1914 – Formicinae: 14 spp., 1 known from PH.

The species found in the Philippines, Anoplolepis gracilipes (F. Smith, 1857), is a pantropical invasive ant, with 11-segmented antennae with extremely long antennal scapes and a slender, constricted mesosoma in dorsal view. It is locally dominant around its nest. This species may nest in the ground or in a tree hollow. Field observations by the present authors indicate that this ant may be native to the Philippines. Upon opening a nest in a tree hollow, we found that muscoid flies immediately hovered over the scampering workers and the brood. In a coconut farm in Candelaria, Quezon Province, the senior author also observed and collected an immature reduviid bug mimicking the erratic movement, color, and size of this ant. These ants may be collected by carefully inspecting tree hollows and rotten logs, and searching for trails of foraging workers on the ground and along branches.

Aphaenogaster

Genus Aphaenogaster Mayr, 1853 – Myrmicinae: 227 spp., at least 2 from PH.

(New record). There are historical specimens of two undescribed species, collected by Chapman and his field crew, of this genus in the ant collection of the Museum of Comparative Zoology, Harvard University. There are also new collections from Mindanao Island. These ants typically nest in rotten logs and the soil under the logs. These ants may be collected by pitfall trapping, breaking into rotten logs and searching for foragers on the ground.

• Aphaenogaster feae

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Bothriomyrmex

Genus Bothriomyrmex Emery, 1869b – Dolichoderinae: 46 spp., at least 2 from PH.

A specimen of an unidentified species, collected from Palawan Island in 1925, is in the ant collection of the Museum of Comparative Zoology, Harvard University. There are specimens of an undetermined species, collected in Laguna Province, Luzon Island, in the Insect Collection of the International Rice Research Institute. An excellent nest series with all the castes of possibly another species was recently collected by Perry Buenavente from Mt. Diwata, Agusan del Sur Province, Mindanao Island. These small ants, superficially similar to Tapinoma, have a petiole with an erect scale; with short, indistinct palps and a propodeum with a short dorsal face and a long declivity. These ants may be collected by sifting leaf litter including the humus layer. Emery (1925b) (out of date); Shattuck (1992c) (generic revision of subfamily).

Figure 2. Full-face and profile images of Philippine ant genera. Amblyopone luzonica (Wheeler and Chapman, 1925) (A, B). Anillomyrma decamera (images reproduced with permission of Dr. Katsuyuki Eguchi) (C, D). Anochetus isolatus Mann, 1919 (E, F).

Calomyrmex

• Calomyrmex species 1

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Calyptomyrmex

Genus Calyptomyrmex Emery, 1887b – Myrmicinae: 25 spp., 2 known from PH.

These small, cryptic, slow-moving ants bear a hard, thick integument with numerous evenly-spaced clavate hairs. They curl up into a ball when disturbed and are found in the leaf litter or on the ground. These ants may be collected by sifting leaf litter on a white sheet, waiting a while and carefully inspecting seed-like objects. After about a minute, these seed-like objects may stretch out and start to walk slowly. Keys: Baroni Urbani (1975a) (Indian subcontinent); Bolton (1981) (Afrotropical); Shattuck (2011) (Southeast Asian revision).

Camponotus

Genus Camponotus Mayr, 1861 – Formicinae: 1,584 spp., at least 28 known from PH.

Camponotus is an extremely large genus in dire need of taxonomic revision. This is a widespread genus in the Philippines. Twenty-eight species are presently known from the Philippines, but this is probably only a fraction of the total. This genus is unusual among formicines in that the usual conspicuous ring of hairs around the acidipore is absent. This genus can be recognized by the placement of the antennal insertions, which are always set back (not adjacent to) from the posterior clypeal border. Camponotus are often medium to large ants; dimorphic or polymorphic workers that forage along trails from their nest in wood. Some species are nocturnal. Mimicry also occurs in this genus. There is a single undetermined minor worker in the Philippine collection of the Bernice P. Bishop Museum that closely mimics ants of the genus Dolichoderus. These ants may be collected by breaking into cavities in live wood or dead branches and searching for foragers on the ground, foliage or in the canopy.

Cardiocondyla

Genus Cardiocondyla Emery, 1869a – Myrmicinae: 69 spp., 5 known from PH.

This genus is known to have both normal winged and worker-like wingless males which fight and exhibit interesting courtship behavior (Heinze et al. 1993, Yamauchi et al. 2005, Mercier et al. 2007). These tiny, ground- and rock-dwelling ants bear the characteristic swollen postpetiole which is wider than long, and much larger than the petiole. Several species are widespread pantropical tramps. There are 5 species known from the Philippines. One species, Cardiocondyla sima Wheeler, 1935, is apparently widespread, reported in Lanao Province, Mindanao Island and in Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These ants may be collected by leaf litter sifting, pitfall trapping and baiting with cookie crumbs on exposed rocks in creeks and rivers and following the forager to its nest in the rock’s crevices. Key to species: Seifert (2003) (Selected species-groups only).

Figure 3. Full-face and profile images of Philippine ant genera. Anoplolepis gracilipes (A, B). Aphaenogaster species PH-02 (C, D). Bothriomyrmex species PH-01 (E, F).

Figure 4. Full-face and profile images of Philippine ant genera. Calyptomyrmex beccarii Emery, 1887 (A, B). Camponotus albocinctus (Ashmead, 1905), major worker (C, D). Cardiocondyla sima, queen (E, F).

Carebara

Genus Carebara Westwood, 1840 – Myrmicinae: 175 spp., at least 3 known from PH.

The Philippine species of this pantropical genus are poorly known. There are specimens of two unidentified species from a transect study at Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). Other species have been collected on Samar Island and in Misamis Occidental Province, Mindanao Island. These are very tiny ants with dimorphic workers. The major workers can possess a pair of tubercles or horns near the posterior margin of the head. They are ground-dwelling. These ants may be collected by sifting soil and leaf litter. Key to species: Fernandez (2004) (New World only), Fernandez (2010) (description of first Philippine species).

Cataulacus

Genus Cataulacus F. Smith, 1853 – Myrmicinae: 69 spp., 3 known from PH.

These robust, hard-bodied ants have a wide, sculptured head and the antennal scrobe passing below the eye. They are known to be arboreal, can glide back to the tree trunk when they fall (Yanoviak et al. 2008) and nest in small twigs or tree trunks. These ants may be collected by searching dead branches still attached to the tree and beating low vegetation over a white sheet. Key to species: Bolton (1974) (World revision).

Centromyrmex

Genus Centromyrmex Mayr, 1866b – Ponerinae: 15 spp., 1 known from PH.

Centromyrmex feae is the only species recorded from the Philippines at present. These small, cryptic ants have a pronotum that is somewhat flattened dorsally, mandibles that sharply curve downward and backward (in side view), and middle tibiae with strong peg-like setae. The workers are weakly polymorphic, with slight differences among nestmates. They are hypogaeic. Some evidence indicates that they are associated with termites. These ants may be collected by sifting leaf litter and soil cores at least 10 cm deep.

Cerapachys

Genus Cerapachys F. Smith, 1857 – Cerapachyinae: 144 spp., 10 known from PH.

These hard-bodied ants have a large, globular petiole and a larger, barrel-shaped postpetiole. They are predators on other ants and some species conduct group raids on other ant nests. During a raid, they steal the larvae and pupae which they sting. These stung prey remain alive for a long time, providing fresh food for the colony. In Mt. Isarog, Bicol Region, Luzon Island, they have been found nesting in twigs, with a larder of pupae of another ant genus, Crematogaster (Alpert and General in prep.). These ants may be collected by breaking open twigs on the ground and in the leaf litter, sifting leaf litter, and searching for columns of raiding workers. Key to species: Brown (1975) (World).

Crematogaster

Genus Crematogaster Lund, 1831 – Myrmicinae: 780 spp., 16 known from PH.

These small ants have a characteristically heart-shaped gaster that can flex over the mesosoma. They are often associated with coccids and aphids, sometimes building carton or soil shelters over these sap-sucking insects. They may be hypogaeic, epigaeic or arboreal. These ants may be collected by breaking open carton or soil that is at the apices of young branches, sifting leaf litter and soil, inspecting plant roots, beating low vegetation over a white sheet, and baiting with peanut butter on tree trunks or cookie crumbs on the ground. Key: Hosoishi and Ogata (2009a) (subgenus Physocrema); Hosoishi and Ogata (2009b) (checklist of Asian species).

Figure 5. Full-face and profile images of Philippine ant genera. Carebara alperti Fernandez, 2010 (A, B). Cataulacus chapmani Bolton, 1974 (C, D). Centromyrmex feae (E, F).

Dacetinops

Genus Dacetinops Brown and Wilson, 1957 – Myrmicinae: 7 spp., 1 known from PH.

(New record). Dr. Herbert Zettel (Natural History Museum, Vienna, Austria) has a single specimen, tentatively identified as Dacetinops cirrosus Taylor, 1985, collected near Calbiga-a River, Mt. Pangasugan, Baybay, Leyte Island. These ants bear spongiform tissue on the ventral surfaces of the petiole, postpetiole, and the first gastral segment. They nest in rotten wood. These ants may be collected by sifting leaf litter and breaking into rotten twigs and wood on the forest floor. Key to species: Brown and Wilson (1957), Taylor (1985) (Papuasian).

Diacamma

Genus Diacamma Mayr, 1862 – Ponerinae: 44 spp., perhaps 8 known from PH.

This genus is long overdue for revision. These ants are easily distinguished from other ponerines by the distinctive costate sculpturing that covers the head, mesosoma, and the petiole, which has 2 dorsal spines. The late Dr. W.L. Brown, Jr. believed that male characters may hold the key to producing stable species boundaries. These large black ants are ground-dwelling or arboreal and hunt singly for prey. Some species also have a unique social structure in which the queen caste is absent and all workers have the potential to mate and lay eggs. Peeters and Higashi (1989) (reproductive dominance behavior).

Dilobocondyla

Key to Dilobocondyla Species of the Philippines

Genus Dilobocondyla Santschi, 1910 – Myrmicinae: 11 spp., at least 3 known from PH.

These small hard-bodied ants have the upper corners of the head drawn into broad points and have a distinctive barrel-shaped petiole. They are known to be arboreal but may forage on the ground as well. An unidentified species is reported from a transect study at Mt. Isarog, Camarines Sur Province, Luzon Island (Alpert and General in prep.). These ants may be collected by beating low vegetation over a white sheet and inspecting dead branches still attached to the tree for nests.

Discothyrea

Genus Discothyrea Roger, 1863 – Proceratiinae: 32 spp., at least several known from PH.

(New record). There are specimens of Discothyrea bryanti (Wheeler, 1917) and D. clavicornis Emery, 1897 (Shattuck, pers. comm.) from a transect study at Mt. Isarog by Joanaviva Caceres-Plopenio in 2006. Another unidentified species was found from a different transect study at Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). Specimens were collected at 550-700 meters above sea level. This is remarkable because few species of this genus are sympatric, much less on the same transect. Two unidentified species were extracted from lowland forest leaf litter berlesate from Samar Island. Another unidentified species was collected from Palawan Island.

Figure 6. Full-face and profile images of Philippine ant genera. Cerapachys rufithorax Wheeler and Chapman, 1925 (A, B). Crematogaster difformis F. Smith, 1857 (C, D). Dacetinops cirrosus (E, F).

These small cryptic ants have an extremely large apical antennal segment and the petiole broadly attached to the gaster. They are known to be predators of arthropod eggs. These ants may be collected by sifting leaf litter, pitfall trapping, and searching for nests under rocks. Brown (1958). Genus Dolichoderus Lund, 1831 – Dolichoderinae: 148 spp, 3 known from PH.

The most common species, Dolichoderus thoracicus (F. Smith, 1860), is the black, hard-bodied ant usually associated with the locally-popular fruit of the lanzones tree (Meliaceae: Lansium domesticum Corr.), and of the makopa tree (Myrtaceae: Syzygium samarangense (Blume) Merrill and Perry) tending the mealybugs (Hemiptera: Sternorrhyncha: Pseudococcidae) that are found in the fruit bunches (DMG, personal observation). This ant, which forages day and night, is also common in urban and highly disturbed areas. These ants are easy to collect, with newly-mated queens even venturing indoors at night.

Echinopla

Genus Echinopla F. Smith, 1857 – Formicinae: 26 spp., 4 known from PH.

Specimens of two unidentified species were collected from low vegetation in a transect study in Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). One of these unidentified species was collected by nocturnal beating of low vegetation. These hairy or fuzzy ants are unique in appearance and have hard bodies and petioles armed with teeth or denticles laterally. These ants may be collected by beating of low vegetation over a white sheet and inspecting of dead branches in the foliage and canopy.

Euprenolepis

Genus Euprenolepis Emery, 1906 – Formicinae: 6 spp., 1 known from PH.

Euprenolepis negrosensis (Wheeler, 1930) is the single species known from the Philippines. It is rarely collected, perhaps because its tiny eyes indicate a subterranean life-habit. These ants have strongly curved mandibles such that the apical tooth is directed posteriolaterally. They forage in the leaf litter and on the ground, possibly for mushrooms, as found by Witte and Maschwitz (2008) in Malaysia. These ants may be collected by sifting leaf litter, pitfall trapping, and possibly by baiting with edible mushrooms. Key to species: LaPolla (2009) (World revision).

Eurhopalothrix

Genus Eurhopalothrix Brown and Kempf, 1960 – Myrmicinae: 36 spp., 3 known from PH.

This genus may be confused with Pyramica, but has more than 6 antennal segments. These small, cryptic, slow-moving ants have clavate hairs all over the body. They forage in the leaf-litter. When disturbed, these ants roll up into a tight ball which makes them even harder to find. After about a minute, they stretch out and start walking slowly. There are specimens of an unidentified species from a transect study at Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). Specimens were collected at 550 meters. These ants may be collected by sifting leaf litter over a white sheet, waiting a while, and carefully inspecting seedlike objects that begin to walk slowly. Keys to species: Brown and Kempf (1960) (World); Taylor (1968, 1980, 1990) (Indo-Australian, Australasian).

Figure 7. Full-face and profile images of Philippine ant genera. Diacamma panayense Wheeler and Chapman, 1925 (A, B). Dilobocondyla chapmani Wheeler, 1924 (C, D). Discothyrea clavicornis (E, F).

Forelophilus

Genus Forelophilus Kutter, 1931 – Formicinae: 3 spp., 2 known from PH.

This genus was previously known only from the type species, Forelophilus overbecki Kutter, 1931, described from Java. Two widespread species are now known from the Philippines, F. stefanschoedli Zettel and Zimmerman, 2007 known from the islands of Luzon, Leyte, and Mindanao, and F. philippinensis Zettel and Zimmerman, 2007 known from the islands of Luzon, Bayagnan, and Mindanao (Zettel and Zimmerman 2007). These ants may be collected by beating low vegetation over a white sheet and hand collecting from foliage.

Gauromyrmex

Genus Gauromyrmex Menozzi, 1933 – Myrmicinae: 2 spp., 1 known from PH.

(New record). Gauromyrmex acanthinus (Karavaiev, 1935) is known from the Philippines. A single identified specimen is deposited in the MCZ Ant Collection. These small ants are generally similar to Vollenhovia but have a propodeum armed with short spines. These ants typically nest under rocks. They can be collected by turning over rocks and other hard objects, sifting leaf litter, and pitfall trapping. Bolton (2003).

Gesomyrmex

Genus Gesomyrmex Mayr, 1868 – Formicinae: 7 spp., 1 known from PH.

Gesomyrmex luzonensis (Wheeler, 1916) is known from the Philippines. This ant is rarely collected but widespread, having been collected in a bamboo grove in Los Baños, Laguna Province, Luzon Island and in a hardwood cavity nest in Cuernos de Negros, Negros Island. All members of this genus are presumed to be nocturnal and arboreal in habitat. These ants have large bean-shaped eyes and mandibles that look like pinking scissors. They are polymorphic, with major, media, and minor workers. Wheeler (1930c) quoted J.W. Chapman’s description of the habits of G. luzonensis. Chapman found that the minors conduct most of the foraging, are attracted to ripe bananas, and are very timid. Key to species: Cole (1949); see also Wheeler (1929a, 1929b, 1930a).

Gnamptogenys

Genus Gnamptogenys Roger, 1863 – Ectatomminae: 134 spp., 11 known from RP.

These hard-bodied ants are specialized predators of other ant species. At least one species, Gnamptogenys menadensis (Mayr, 1887), has mostly fertile laying workers instead of a queen (Gobin et al. 1998, Gobin et al. 1999). They may be ground-dwelling or arboreal, diurnal or nocturnal. These ants may be collected by breaking open rotten wood on the ground, inspecting cavities in living trees and dead branches, sifting leaf litter, and pitfall trapping. Key to species: Lattke (2004) (SE Asia and Australasia).

Figure 8. Full-face and profile images of Philippine ant genera. Dolichoderus thoracicus (A, B). Echinopla pallipes F. Smith, 1857 (C, D). Euprenolepis species PH-01 (E, F).

Figure 9. Full-face and profile images of Philippine ant genera. Eurhopalothrix philippina Brown and Kempf, 1960 (A, B). Forelophilus species PH-01 (C, D). Gauromyrmex acanthina (E, F).

Harpegnathos

Genus Harpegnathos Jerdon, 1851 – Ponerinae: 11 spp., 4 known from PH.

These large cryptic ants bear characteristic long pliers-like mandibles and large eyes. They are usually ground-dwelling and hunt prey such as crickets in the leaf litter. A specimen of an unidentified arboreal species was collected on Mt. Isarog, Bicol Region, Luzon Island (DMG, unpublished notes). These ants may be collected by sifting leaf litter, searching on the ground, and pitfall trapping.

Hypoponera

Genus Hypoponera Santschi, 1938 – Ponerinae: 171 spp., 5 known from PH.

The taxonomy of this large pantropical genus is currently chaotic. These small, cryptic ants are very similar to Pachycondyla, but lack a second simple spur on the hind tibia. They have a simple ventral process of the petiole, without a fenestra or posterior angles as in Ponera. These ants make small colonies in soil, rotten wood, and leaf litter. These ants may be collected by sifting leaf litter, beating low vegetation, pitfall trapping, and searching in rotten wood. There is no modern key to species, however, Barry Bolton is currently revising this genus (pers. comm.).

Iridomyrmex

Genus Iridomyrmex Mayr, 1862 – Dolichoderinae: 82 spp., at least 1 known from PH.

These fast-moving ants may be confused with Dolichoderus but have a thinner cuticle and bear the characteristic wavy anterior clypeal margin. A specimen of Iridomyrmex anceps (Roger, 1863) was recently collected from Marinduque Island by Joanaviva Caceres-Plopenio (Ward, pers. comm). There is a specimen of one unidentified species, collected from Baguio, Benguet Province, Luzon Island, in the MCZ ant collection. Other unidentified specimens have been collected from the provinces of Pangasinan, Isabela, and Camarines Sur on Luzon Island, and the islands of Camiguin and Semirara. They are ground-dwelling and usually form large colonies. These ants may be collected by sifting leaf litter, pitfall trapping, and baiting on the ground. Generic review: Shattuck (1992a).

Lepisiota

Genus Lepisiota Santschi, 1926 – Formicinae: 131 spp., 2 known from PH.

These small, yellow or brown arboreal ants have a distinct angulate propodeum, and are rarely collected. They have been collected in the islands of Luzon, Negros, and Palawan. These ants may be collected by beating low vegetation over a white sheet and inspecting cavities in living wood and dead branches in the canopy.

Leptanilla

Genus Leptanilla Emery, 1870 – Leptanillinae: 43 spp., 2 known from PH.

Leptanilla astylina Petersen, 1968 is known from the Philippines (Palawan). These extremely small, blind, hypogaeic ants have a slim, elongated body and a swollen postpetiole. They are extremely difficult to collect, perhaps because they live deep in the soil. These ants may be collected by sifting soil cores up to 30 cm deep and underground baiting. Key to species: Baroni Urbani (1977b) (World), Ogata et al. (1995).

Described from an alate male reproductive, collected in southern Palawan Island, is Leptanilla copiosa Petersen, 1968 (previously Noonilla copiosa). Ogata et al. (1995) examined a male specimen from Misamis Oriental Province, Mindanao Island and considered this genus incerta sedis in the subfamily. Bolton (2003) excluded the genus from Formicidae. However, Marek Borowiec (pers. comm.), who is currently studying the subfamily, is convinced that Noonilla belongs to Leptanillinae. These ants may be collected by sifting soil cores up to 30 cm deep, underground baiting, and Malaise trapping for alates. Ogata et al. 1995, Bolton (2003) (synopsis of Formicidae).

Figure 10. Full-face and profile images of Philippine ant genera. Gesomyrmex luzonensis, major worker (A, B). Gnamptogenys chapmani Brown, 1958 (C, D). Harpegnathos venator Donisthorpe, 1937 (E, F).

Figure 11. Full-face and profile images of Philippine ant genera. Hypoponera confinis (Roger, 1860) (A, B). Iridomyrmex species PH-01 (C, D). Lepisiota chapmani Wheeler, 1935 (E, F).

Figure 14. Full-face and profile images of Philippine ant genera. Mayriella transfuga (A, B). Meranoplus species PH-01 (C, D). Metapone gracilis Wheeler, 1935 (E, F).

Figure 15. Full-face and profile images of Philippine ant genera. Monomorium species PH-01 (A, B). Myopias lobosa Willey and Brown, 1983 (C, D). Myopopone castanea (E, F).

Figure 16. Full-face and profile images of Philippine ant genera. Myrmecina species PH-01 (A, B). Myrmicaria brunnea Saunders, 1842 (C, D). Myrmoteras wlliamsi Wheeler, 1919 (E, F).

Figure 17. Full-face and profile images of Philippine ant genera. Mystrium camillae Emery, 1889 (A, B). Noonilla copiosa (line drawings reprinted from Entomologiske Meddelelser, with permission) (C, D). Nylanderia species PH-01 (E, F).

Leptogenys

Genus Leptogenys Roger, 1861 – Ponerinae: 248 spp., 11 known from PH.

These long-bodied and slender ants bear the characteristic clypeus extending forward to form a rounded triangle. They also exhibit army ant-like behavior with large raiding columns and are known to prey on termites. They usually have worker-like, or ergatoid, queens and nest in rotten wood, under rocks, and in the ground. These ants may be collected by sifting leaf litter, pitfall trapping, and searching for raiding columns. Key to species: Wilson (1958) (Melanesian and New Caledonia).

Leptomyrmex

Genus Leptomyrmex Mayr, 1862 – Dolichoderinae: 41 spp., 1 known from PH.

This genus occurs mostly in Australia, New Guinea, New Caledonia, and Aru Island, Indonesia. Leptomyrmex is unique in the subfamily in having an elongated head, mesosoma, and legs. In addition, most of the queens are wingless. A single specimen of Leptomyrmex fragilis (F. Smith, 1859) has been found in the Bernice P. Bishop Museum collection. The specimen was collected by L. W. Quate in 1959 in San Francisco, Agusan del Sur Province, Mindanao Island. Lucky and Ward (2010) consider this specimen a doubtful record, however it is validated by the many legitimate records of other insects in the Quate collection. These ants may be collected by sifting leaf litter, pitfall trapping, baiting on the ground, and searching for nests at the base of trees and in rotten wood. Key to species: Lucky and Ward (2010). Generic review: Shattuck (1992c).

Liomyrmex

Genus Liomyrmex Mayr, 1865 – Myrmicinae: 1 sp., 1 known from PH.

This is a monotypic tropical genus, with only one widespread species. Liomyrmex gestroi Emery, 1887 is known from throughout Southeast Asia. In the Philippines, it has been recorded from the islands of Luzon, Mindanao and Negros. These tiny, blind ants have a smooth almost hairless body and bear large ventral processes on the petiole and postpetiole. They are assumed to be a kleptoparasite of mound-building termites, having been found inside live termite mounds and in forest leaf litter with termites. These ants may be collected by sifting leaf litter, pitfall trapping, and breaking into termite mounds to find nests. Rigato and Bolton (2001) (World revision).

Lophomyrmex

Genus Lophomyrmex Emery, 1892 – Myrmicinae: 12 spp., 1 known from PH.

(New record). Specimens of Lophomyrmex bedoti Emery, 1893 were recently collected by general collecting in Palawan. These monomorphic ants are hunter/scavengers, preying on various arthropods on the forest floor. They nest near or at the base of trees and form persistent soil-walled trails (similar to the trails of Pheidologeton ants), which may also run underground. These ants may be collected by baiting with sugar or protein bait and searching for nests at the base of trees and for conspicuous trails on the forest floor. Key to species: Rigato (1994) World revision.

Lordomyrma

Genus Lordomyrma Emery, 1897a – Myrmicinae: 20 spp., at least 4 known from PH.

(New record). There are specimens of two unidentified species from Mt. Isarog, Bicol Region, Luzon Island. A specimen of another unidentified species was collected in a transect study in Bulacan Province, Luzon Island. Another unidentified species from Mt. Maquiling (R. Morse, leg. 1968) is present in the Ant Collection of the MCZ at Harvard University. These small monomorphic ants are variable in morphology but usually have a prominently convex pronotum. These ants forage in the foliage and on the ground. Little is known of their biology. These ants may be collected by beating low vegetation, sifting leaf litter, flipping rocks and breaking into rotten wood for nests. Keys to species: Wheeler (1919b), Donisthorpe (1941a) (both outdated).

Mayriella

Genus Mayriella Forel, 1902c – Myrmicinae: 7 spp, 1 known from PH.

The species known from the Philippines, Mayriella transfuga Baroni Urbani, 1977, is wide-ranging in the Asian tropics and was originally described from Nepal. These are tiny, hard-bodied, ground-foraging ants with a clypeus that extends as two lobes over the mandibles. The head and mesosoma are heavily sculptured while the gaster is smooth. They nest in rotten wood and under stones. These ants may be collected by sifting leaf litter, pitfall trapping, and flipping stones and breaking into rotten wood for nests. Key to species: Shattuck and Barnett (2007) (World revision).

Meranoplus

Genus Meranoplus F. Smith, 1853 – Myrmicinae: 62 spp., 2 known from PH.

Only one species, Meranoplus biliran Schödl, 1998, is known from Biliran Island. There are specimens of another, possibly undescribed, species from the nearby island of Samar. These slow-moving ants have the characteristic shield-like upper surface of the mesosoma, which is actually an extension of the pronotum. When disturbed, they curl up and remain motionless, similar to Eurhopalothrix and Calyptomyrmex ants. They are ground-dwelling generalist scavengers or seed predators and forage on the ground an in the foliage. These ants may be collected by beating low vegetation, sifting leaf litter, pitfall trapping, baiting with tuna or honey on trees, and searching for a ring of seed hulls around the nests. Key to species: Schödl (1998) (Oriental).

Figure 12. Full-face and profile images of Philippine ant genera. Leptanilla astylina (line drawing reprinted from Entomologiske Meddelelser, with permission) (A). Leptogenys maxillosa (F. Smith, 1858) (B, C). Leptomyrmex fragilis (D, E).

Figure 13. Full-face and profile images of Philippine ant genera. Liomyrmex gestroi (A, B). Lophomyrmex bedoti (C, D). Lordomyrma species PH-01 (E, F).

Metapone

Genus Metapone Forel, 1911c – Myrmicinae; 17 spp., 2 known from PH.

The two Philippine species are known only from winged reproductives. An unidentified species is present in the Bernice P. Bishop Museum ant collection. These hard-bodied cryptic ants are commonly mistaken for ponerine ants because of their large, broadly attached postpetiole. These ants have deep antennal scrobes and the clypeus projects forward as a square lobe. They are often found feeding on hardwood termites in dead logs and are very rarely collected. These ants may be collected by breaking into the heartwood of hardwood logs that are suspended off the ground. Key to species: Wheeler, (1919c) (World), Alpert (2007) (Madagascar).

Monomorium

Genus Monomorium Mayr, 1855 – Myrmicinae: 399 spp., 5 known from PH.

These tiny, smooth and slender ants have a single strong seta on the anterior clypeal margin. This genus includes worldwide invasive species such as M. pharaonis (Linnaeus, 1758) and M. floricola Forel, 1910. These ants are typically opportunistic predators and scavengers. Their ground-nests are usually marked by a small crater of excavated soil. These ants may be collected by sifting leaf litter and pitfall trapping and searching for their nests on the ground. Key to species: Heterick (2001) (Australian).

Myopias

Genus Myopias Roger, 1861 – Ponerinae: 36 spp., 5 known from PH.

These cryptic ants have eyes that are very close to the base of the mandibles, and usually have a clypeal extension visible in the large gap between the mandibles. They nest in rotten twigs and logs and forage in the leaf litter and on the ground. They are known to prey on springtails, millipedes, and even other ants. These ants may be collected by sifting leaf litter and breaking into rotten wood. Willey and Brown (1983) (Australasian).

Myopopone

Genus Myopopone Roger, 1861 – Amblyoponinae: 1 spp, 1 known from PH.

The sole species, Myopopone castanea (F. Smith 1860), is widespread in the Philippines. These large-headed ants have the characteristic flattening of the antennal flagella. They nest in Zorapteran-stage rotten wood, apparently preying on beetle larvae. These ants may be collected by breaking open rotten logs that you can dig into with a garden trowel. Brown (1960) (World revision).

Myrmecina

Genus Myrmecina Curtis, 1829 – Myrmicinae: 37 spp., at least 3 known from PH.

This genus is widespread throughout the Philippines. There are specimens of several unidentified species from different transect studies on Mt. Isarog, Bicol Region, Luzon Island (Samson et al. 1997, Caceres in prep., Alpert and General in prep.). One of these species seems to mimic Pristomyrmex ants. There are also specimens of another species from transect studies in the provinces of Bulacan and Nueva Vizcaya, Luzon Island. Specimens of yet another unidentified species were collected in Palawan Island. These hard-bodied ants have the characteristic ventral ridge of the head, running from the back of the head to the base of the mandibles, and the distinctive barrel-shaped petiole. They nest in twigs and rotten wood or under rocks and forage in the leaf litter. These ants may be collected by sifting leaf litter, pitfall trapping, and flipping over rocks.

Myrmicaria

Genus Myrmicaria Saunders, 1842 – Myrmicinae: 67 spp., 2 known from PH.

These relatively large ants bear the characteristic 7-segmented antennae and long anterior peduncle of the petiole. They form conspicuous columns, forage on the ground and in the foliage, and are locally dominant where they occur. These ants are fairly easy to collect by baiting with tuna or honey, pitfall trapping, beating low vegetation, and hand collecting. Santschi (1925) (African; out of date).

Myrmoteras

Key to Philippine species of Myrmoteras

• 

  Myrmoteras williamsi

• 
• 
• 

Genus Myrmoteras Forel, 1893 – Formicinae: 32 spp., at least 4 known from PH.

• Myrmoteras glabrum
• Myrmoteras insulcatum
• Myrmoteras mcarthuri
• Myrmoteras williamsi

These small trap-jawed ants have long mandibles that look like ripsaws and have very large eyes. They nest and hunt soft-bodied arthropods in the leaf litter. These ants may be collected by inspecting dead leaves that stick together then sifting the leaf litter. Keys to species: Moffett (1985) (World revision), Agosti (1992) (World revision), Zettel and Sorger (2011) Philippine species.

Zettel and Sorger (2011) - The genus has two subgenera, Myrmoteras s.str. and Myagroteras Moffett, 1985. They are probably monophyletic entities (Moffett 1985). Only Myagroteras is known east of Huxley’s Line (Philippines, Sulawesi, Lombok), and two Philippine-endemic species have been described in the past: Myrmoteras (Myagroteras) williamsi Wheeler, 1919 and Myrmoteras (Myagroteras) insulcatum Moffett, 1985. Knowledge on Myrmoteras in the Philippines is extremely fragmentary, and much more specific field work with leaf litter sampling is required in order to provide a more complete picture than we can present now.

Mystrium

Genus Mystrium Roger, 1862 – Amblyoponinae: 9 spp., 1 known from PH.

These cryptic ants have long, linear mandibles, inserted at the sides of the head, with blunt ends and a snaggle-tooth arrangement. These ground-foraging ants usually have soil particles stuck to their body, providing perfect camouflage. When disturbed, they lie motionless. They nest under rocks and in rotten wood. These ants may be collected by sifting leaf litter, flipping over rocks, and breaking into rotten logs. Brown (1960) (World revision), Bihn and Verhaagh (2007) (Indo-Australian, with tabular key).

Nylanderia

Genus Nylanderia Emery, 1906 – Formicinae: 133 spp., at least 3 spp. known from PH.

This genus was recently revived from synonymy by LaPolla et al. 2010. There are at least 3 unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.), including a mimic of Tapinoma melanocephalum (Fabricius, 1793). These ants typically forage in the foliage, on the ground and in the leaf litter. They feed on honeydew from scale insects and aphids, and scavenge dead insects. These ants may be collected by beating of low vegetation, sifting leaf litter, pitfall trapping, and hand collecting. LaPolla et al. 2010 (Generic key to Prenolepis genus-group).

Ochetellus

Genus Ochetellus Shattuck, 1992a – Dolichoderinae: 10 spp., 1 known from PH.

The only species known from the Philippines, Ochetellus glaber (Mayr, 1862), is widespread in the archipelago. These fast-moving ants have a concave anterior clypeal margin and concave propodeum. They may be arboreal or ground-foraging, sometimes forming conspicuous columns. These ants may be collected by beating of low vegetation, sifting leaf litter, and pitfall trapping. Shattuck (1992a) (generic revision).

Odontomachus

Genus Odontomachus Latreille, 1804 – Ponerinae: 65 spp., 11 known from PH.

Key to Philippine Odontomachus

These large, big-headed, trap-jawed ants are commonly referred to as "hantik" in many Philippine dialects. They are aggressive, locally dominant and ground-dwelling but may also climb the foliage to hunt for prey. One species, Odontomachus malignus F. Smith, 1859, is known to nest in rock crevices that are regularly inundated during high tide. At low tide, they emerge and forage for animals trapped in the intertidal zone. These ants may be collected by baiting with tuna, beating of low vegetation, pitfall trapping, and hand collecting. Keys to species: Brown (1976) (World), Sorger and Zettel (2011) (Philippine species).

Sorger and Zettel (2011) - Although Odontomachus are very conspicuous ants, little attention has been paid to their taxonomy since Brown (1976) revision. As we have mentioned previously, the Philippines harbour two sets of Odontomachus species, one of which is generally widespread in distribution and the other of which, the one we discuss here in more detail, includes more range-restricted species. Regarding the latter set, what might be called the Philippine O. infandus group species, Brown's (1976) concept of a widely distributed, highly variable species, O. infandus, turned out to be wrong; perhaps simply because it was blurred by incorrect label data. New and correctly labelled samples show a pattern of distinct, sympatric and allopatric species, each with a comparatively stable character set and endemic on one or a few islands. Sympatry is observed only on the island of Luzon (four species), allopatry on several other islands. In other words, there appears to be a radiation of native Odontomachus species both among and within islands. Ranges of the more widely distributed species agree relatively well with patterns found in other terrestrial or limnic organisms and are largely effected by the areas covered by the large Pleistocene islands (e.g., Ong et al. 2002 and references cited therein).

The present study includes two unnamed species. Species 1 from southern Luzon is somewhat problematic as it shows affinities to the sympatric species, O. infandus and O. banksi (in one instance even sharing the same habitat with the latter), and similarities with the allopatric O. alius. Species 2, although only based on one individual, must be regarded as an undescribed species, but it seems advisable to see more specimens before naming it.

A key next step is to study the molecular phylogenetic relationships of species and isolated populations of the Philippine O. infandus-group species. The archipelago is among the earth's most important biodiversity hot-spots (Mitter-Meier et al. 2005, Catibog-Sinha and Heaney 2006) and a “laboratory of evolution”, many thousand times more diverse than the famous Galapagos Islands. Odontomachus species, with their conspicuous and consequential feeding morphology have the potential to serve as living laboratory for understanding evolution of this diverse region, not quite Darwin's finches, but perhaps Brown's trap-jaws.

There is a lack of taxonomic knowledge of the sexuals of most Odontomachus species. Males are presently not identifiable at all. Excavations of complete nests (e.g., see Tschinkel 2011) would help, also for learning about size and development of Odontomachus colonies. Focused studies of these species in general would be rewarding. It is perhaps worth noting in this regard that the collections by the second author and co-fellows are by-products of his project on aquatic Hemiptera (Gapud and Zettel, 1999) rather than focussed ant research.

Another area for promising research might be to consider the biology of the enigmatic species, O. malignus. There are only a few anecdotal notes on its life (Wilson 1959, Brown 1976, Olsen 2009), but it is well known that ants in intertidal zones can display fascinating adaptations to their unfavourable habitats (Nielsen 2011).

Finally, we would be remiss if we did not mention the conservation implications of our findings. Ants do have a potential importance for conservation biology (e.g., Kautz and Moreau 2011 and references therein), particularly in tropical countries. Sadly, destruction of forests in the Philippines is still ongoing, despite the present low level of natural forest cover (Ong et al. 2002, Catibog-Sinha and Heaney 2006). Our results suggest that based on morphology, ants of the Philippines may include more endemic species than currently recognized. The number seems likely to increase further with molecular work. Forest-inhabiting, endemic ants are strongly affected and threatened by extinction and range-restricted taxa are more threatened than those that are widespread. This danger is perhaps most acute for species on the Visayas Islands (O. philippinus and O. alius), where forests have been diminished to a few remnant spots (e.g., on Panay, Negros, Siquijor, Cebu, Bohol; see Ong et al. 2002), as well as for the very locally distributed O. scifictus on Camiguin. The more we see the unique elements of local species and forms in Odontomachus, as well as more generally, the more we are aware of their threat and, similarly, just how little we know of what is being lost.

Odontoponera

Genus Odontoponera Mayr, 1862 – Ponerinae: 1 sp., 1 known from PH.

There is one valid species, Odontoponera denticulata (F. Smith, 1858), which is widespread in the Philippines. These moderately large, hard-bodied ants have large blunt teeth on the sides of the pronotum and a crenulate anterior clypeal margin. They nest under bare ground and hunt singly. The nest entrance is a simple hole just large enough for one worker to pass through. These ants may be collected by sifting leaf litter, pitfall trapping, and hand collecting. Wheeler and Chapman (1925), Creighton (1929) (out of date), Yamane (2009).

Oecophylla

Genus Oecophylla F. Smith, 1860b – Formicinae: 2 spp., 1 known from PH.

Oecophylla smaragdina (Fabricius, 1775) is widespread in the Philippines. These large yellow-green ants weave silken nests among the leaves of mango (Anacardiaceae: Mangifera indica L.) and other trees. They are aggressive and dominant where they occur, effectively excluding other ant species in the trees and on the ground as well. A single colony may construct many nests among several trees, with the outer nests serving as defensive bivouacs for older workers and the inner nests containing the queen and brood. Ilokanos, many Southeast Asian tribes, and south Chinese tribes harvest the pupae

Figure 18. Full-face and profile images of Philippine ant genera. Ochetellus glaber (A, B). Odontomachus simillimus F. Smith, 1858 (C, D). Odontoponera denticulata (E, F).

as a delicacy. This species is usually found at elevations below 500 meters. These ants may be collected by searching for nests in the canopy and hand collecting from the tree trunks.

Figure 19. Full-face and profile images of Philippine ant genera. Oecophylla smaragdina (A, B). Overbeckia subclavata Viehmeyer, 1916 (C, D). Pachycondyla claudata (Menozzi, 1926) (E, F).

Overbeckia

Genus Overbeckia Viehmeyer, 1916 – Formicinae: 1 sp., 1 known from PH.

(New record). There is only one valid species in this genus, Overbeckia subclavata. This ant has an antenna that gradually widens to a relatively broad terminal segment, and has a mesosoma that is flat and pinched at the metapleural spiracles. It was previously known only from Singapore, but a specimen in the Museum of Comparative Zoology is labeled "Bu of Sci, PI" and was collected by a certain M. Ramos (probably Maximo Ramos, who collected botanical specimens for the Bureau of Science from 1904 to 1932) (Van Steenis-Kruseman, 2006). “PI” is the abbreviation for Philippine Islands, the old name used during the American colonial period. O. subclavata is a very rare ant, and its rediscovery will be an important event in Philippine myrmecology. This ant species is assumed to be arboreal because of its relatively large compound eyes. These ants may be collected by beating of low vegetation.

Pachycondyla

Genus Pachycondyla F. Smith, 1858 – Ponerinae: 289 spp., 13 known from PH.

This large genus is also in dire need of taxonomic revision. These are small or large ants with two tibial spurs, a large pectinate spur behind a small simple one. They are abundant in the leaf litter and on the ground, hunting singly. These ants may be collected by sifting leaf litter, pitfall trapping, and baiting with honey or tuna bait. The genus Cryptopone was recently synonymized by MacKay and MacKay (2010).

Paraparatrechina

Genus Paraparatrechina Donisthorpe, 1947 – Formicinae: 28 spp, at least 3 known from PH.

This genus was recently revived from synonymy by LaPolla et al. 2010, who split the genus Paratrechina into 3 separate genera. Among these 3 genera, only Paraparatrechina ants have a pair of erect setae on the propodeum and erect setae on the pronotum and mesonotum arranged in neat pairs. P. iridescens (Donisthorpe, 1942) and an unidentified species are known from several locations on Luzon Island. These ants may be collected by sifting leaf litter, pitfall trapping, and baiting with honey or tuna bait. LaPolla et al. 2010 (Generic key to Prenolepis genus-group).

Paratopula

Genus Paratopula Wheeler, 1919a – Myrmicinae: 10 spp., 3 known from PH.

There are ten valid species in this Oriental and Indo-Australian genus, with three species known from the Philippines. These large arboreal ants have a pronotum with a flat dorsal outline. They are rarely collected. These ants may be collected by beating low vegetation over a white sheet and inspecting tree hollows and dead branches in the canopy. Key to species: Bolton (1988) (World).

Figure 20. Full-face and profile images of Philippine ant genera. Paraparatrechina iridescens (A, B). Paratopula macta Bolton, 1988 (C, D). Paratrechina longicornis (Latreille, 1802) (E, F).

Figure 21. Full-face and profile images of Philippine ant genera. Pheidole hortensis Forel, 1913, major worker (A, B). Pheidologeton maccus Wheeler, 1929, major worker (C, D). Philidris species PH-01 (E, F).

Paratrechina

Genus Paratrechina Motschoulsky, 1863 – Formicinae: 1 sp., 1 known from PH.

As revised by LaPolla et al. (2010), this genus now contains only 1 species, the invasive species, Paratrechina longicornis, widespread throughout the Philippines. These ants readily invade households, farms and other highly-disturbed areas, recruiting large numbers of nestmates to scavenge dead insects and small animals and even table scraps. These ants may be collected by sifting leaf litter, pitfall trapping, and baiting with honey or tuna bait. LaPolla et al. 2010 (Generic key to Prenolepis genus-group).

Pheidole

Genus Pheidole Westwood, 1839 – Myrmicinae: 1,121 spp., at least 19 known from PH.

There are specimens of several unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). There are also unidentified species from transect studies in Isabela Province, Luzon Island and the islands of Samar and Mindanao. These tiny to small ants have the following character states: dimorphic, with large-headed major workers possessing usually edentate mandibles; pronotum strongly humped; and antennal club usually 3-segmented. They are ground-dwelling and forage on the ground and in the leaf litter. These ants may be collected by sifting leaf litter, pitfall trapping, and baiting with cookie crumbs. Keys to species: Eguchi (2001a) (Asian), (2001b) (Bornean), (2003, 2004).

Pheidologeton

Genus Pheidologeton Mayr, 1862 – Myrmicinae: 49 spp., 5 known from PH.

This is another genus which needs taxonomic revision. These tiny to small ants form conspicuous raiding columns, often protected by low walls of soil. They are extremely polymorphic with a continuous series of intermediates. Superficially similar to Pheidole, they can be distinguished by their 2-segmented antennal club and their polymorphism. And unlike Pheidole, the major and supermajor workers join the foraging column. They are ground-dwelling. These ants may be collected by sifting leaf litter, pitfall trapping, baiting with honey or tuna bait, and hand collecting. Ettershank (1966).

Philidris

Genus Philidris Shattuck, 1992a – Dolichoderinae: 16 spp., at least 1 known from PH.

Philidris myrmecodiae (Emery, 1887) was reported from Mt. Isarog, Camarines Sur, Luzon Island by Samson et al. (1997). There are specimens of an unidentified species, collected from Zamboanga, Mindanao Island, in the MCZ Ant Collection. There are also specimens from a transect study conducted in Isabela, Luzon Island. These small polymorphic ants are superficially similar to Iridomyrmex but have their eyes very low on the head. They nest in rotten wood above the ground and in swollen specialized plant structures called domatia, which they defend vigorously (Shattuck and Barnett 2010). These ants may be collected by beating low vegetation over a white sheet and inspecting the swollen bases of epiphytic plants. Shattuck (1992a, 1992b).

Plagiolepis

Genus Plagiolepis Mayr, 1861 – Formicinae: 86 spp., at least 2 known from PH.

(New record). There are specimens of an unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.) and of other species from transect studies conducted in the provinces of Isabela and Nueva Vizcaya, Luzon Island. These tiny, cryptic ants have 11-segmented antennae and long palps. They nest in the ground under rocks and in rotten wood and forage in the leaf litter, on the ground, and in the foliage. These ants may be collected by sifting leaf litter, pitfall trapping, and beating low vegetation over a white sheet. Brown (1973).

Platythyrea

Genus Platythyrea Roger, 1863 – Ponerinae: 46 spp., 3 known from PH.

There are 46 valid species, including three from amber, in this cosmotropical genus. There are three valid species known from the Philippines. These ants have a shagreened or dull body surface and two pectinate tibial spurs on the hind leg. They may be ground-dwelling or nesting in trees, and are often found hunting individually. These ants may be collected by sifting leaf litter, pitfall trapping, and hand collecting on tree trunks. Key to species: Brown (1975) (World).

Polyrhachis

Genus Polyrhachis F. Smith, 1857 – Formicinae: 603 spp., at least 66 known from PH.

This is the largest ant genus in the Philippines. There are specimens of several unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These small to large ants have spines on the pronotum, mesonotum, propodeum, or petiole or a combination of locations. They may nest in the ground, rotten logs or tree hollows. Some species are also known as weaver ants because they build nests in the foliage from larval silk and chewed-up plant fibers. These ants may be collected by beating low vegetation over a white sheet, pitfall trapping, and inspecting tree hollows and dead branches in the canopy. Keys: Hung (1967b) (World, subgenera); Kohout (1987) (sexspinosa species-group), Dorow (1995), Dorow and Kohout (1995), Kohout (1998), Kohout (2006a) (cryptoceroides species-group), (2006b) (parabiotica species-group). Natural history: Dorow et al. 1990.

Ponera

Genus Ponera Latreille, 1804 – Ponerinae: 55 spp., 2 known from PH.

These tiny cryptic ants have a fenestra or translucent window in the ventral petiolar process. They forage in the leaf litter and on the ground and nest in rotten wood and under rocks. These ants may be collected by sifting leaf litter, pitfall trapping, breaking into rotten wood, and flipping over rocks. Key to species: Taylor (1967) (World, revision).

Figure 22. Full-face and profile images of Philippine ant genera. Plagiolepis species PH-01 (A, B). Platythyrea parallela (F. Smith, 1859) (C, D). Polyrhachis cyaniventris F. Smith, 1858 (E, F).

Figure 23. Full-face and profile images of Philippine ant genera. Ponera oreas (Wheeler, 1933) (A, B). Prenolepis species PH-01 (C, D). Prionopelta kraepelini Forel, 1905 (E, F).

Prenolepis

Genus Prenolepis Mayr, 1861 – Formicinae: 25 spp., at least 1 known from PH.

There are specimens of an unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These ants have the following character states: very long antennal scapes, at least half the length extending beyond the back of the head; mandibles not strongly curved, so that the apical tooth points to the side; and mesosoma elongated and constricted at midlength. They are ground-foraging. These ants may be collected by sifting leaf litter and pitfall trapping. LaPolla et al. 2010 (Generic key to Prenolepis genus-group).

Prionopelta

Genus Prionopelta Mayr, 1866a – Amblyoponinae: 13 spp., 1 known from PH.

Prionopelta kraepelini was collected from a botanical transect study on Samar Island. This species is widespread, also known from several locations on the islands of Luzon and Negros. Specimens are deposited in the Philippine National Museum in Manila, the UPLB-MNH, and the MCZ Ant Collection. These tiny, cryptic ants have mandibles with only 3 teeth. They forage in leaf litter and on the ground. These ants may be collected by sifting leaf litter and pitfall trapping. Keys to species: Brown (1960) (Indo-Australian, Neotropical), Shattuck (2008a) (Indo-Pacific).

Pristomyrmex

Key to Pristomyrmex of the Philippines

Genus Pristomyrmex Mayr, 1866b –Myrmicinae: 56 spp., 16 known from PH.

Zettel (2006) - At present, 55 species of Pristomyrmex are described (Wang 2003, and this paper). Seventeen species are recorded from the Philippines (= 30.9 % of world fauna). Of these seventeen species, nine species seem to be endemic to the Philippines (= 16.4 % or one sixth of world fauna). The question, whether species are regionally endemic or not, needs to be answered with caution due to the relatively small number of collections.

General and Alpert (2012) - Most species records are from Greater Mindanao (10 species), Greater Luzon (7 species), and Greater Negros-Panay (5 species), probably because of the more intensive collecting activities and more suitable habitats on the larger islands. There are specimens of an unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These small attractive ants have the following character states: mandibles that are twisted so that the edges oppose each other; and exposed antennal sockets. They nest in rotten wood on the ground or under rocks and forage in the leaf litter and on the ground. These ants may be collected by sifting leaf litter, pitfall trapping, breaking into rotten logs and flipping over rocks. Keys to species: Wang (2003) (World revision), Zettel (2006, 2007) (Philippine species).

Probolomyrmex

Genus Probolomyrmex Mayr, 1901 – Proceratiinae: 16 spp., at least 2 known from PH.

One species is known from Negros Island, Probolomyrmex dammermani Wheeler, 1928. There are specimens of an unidentified species collected by Joanaviva Caceres-Plopenio from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Eguchi, pers. comm.; DMG, unpublished notes). These tiny, cryptic ants have no eyes and bear their thick antennae on a shelf projecting forward from the head. They forage in the leaf litter and presumably also in the soil. These are among the rarest ants in the world. These ants may be collected by sifting leaf litter. Status of genus: Brown (1975). Keys to species: Taylor (1965), Eguchi et al. 2006.

Figure 24. Full-face and profile images of Philippine ant genera. Pristomyrmex bicolor Emery, 1900 (A, B). Probolomyrmex dammermani (C, D). Proceratium papuanum Emery, 1897 (E, F).

Figure 25. Full-face and profile images of Philippine ant genera. Pseudolasius typhlops Wheeler, 1935, major worker (A, B). Pyramica pedunculata (Brown, 1953) (C, D). Recurvidris species PH-01 (E, F).

Proceratium

Genus Proceratium Roger, 1863 – Proceratiinae: 80 spp., 1 known from PH.

The species known from the Philippines, Proceratium papuanum was collected from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These tiny ants have the following character states: apical segment of the antenna not extremely large or bulbous; and the petiole narrowly attached to the gaster. They forage in the leaf litter and on the ground. These ants may be collected by sifting leaf litter. Key to species: Baroni Urbani and de Andrade (2003) (World revision, including fossils).

Pseudolasius

Genus Pseudolasius Emery, 1887b – Formicinae: 64 spp., at least 1 known from PH.

This genus can be found in all the Old World tropics except Madagascar. Pseudolasius typhlops is known only from the Philippines. There are historical specimens of a different, unidentified species in the ant collection of the UPLB-MNH, and at least one unidentified species reported by Samson et al. 1997. Other unidentified species were recently collected from other parts of Luzon and from Mindanao Island. These are tiny to small, yellow polymorphic ants. They nest in rotten wood on the ground or underground, where they tend root-feeding coccids. These ants may be collected by sifting leaf litter and soil cores and underground baiting.

Rhopalomastix

Genus Rhopalomastix Forel, 1900a – Myrmicinae; 7 spp., 1 known from PH.

(New record). There are specimens of an unidentified species from Dumaguete, Negros Island in the MCZ Ant Collection. These ants have a longitudinally rugose head and mesosoma, and a postpetiole broadly attached to the gaster. They are presumed to nest and forage in tunnels under bark. These ants are rarely collected. These ants may be collected by searching under bark. Wheeler (1929c) (Review of genus).

Rhoptromyrmex

Genus Rhoptromyrmex Mayr, 1901 – Myrmicinae: 10 spp., 1 known from PH.

There is one specimen, from “Boguio” [=Baguio City, Benguet Province, Luzon Island], of Rhoptromyrmex wroughtonii Forel, 1902 in the MCZ Ant Collection of Harvard University. This species was recently found to be abundant by Joanaviva Caceres-Plopenio during a transect study of an abandoned farm on Mt. Isarog, Camarines Sur, Luzon Island and was also collected by Perry Buenavente from a transect study on Mt. Palali, Nueva Vizcaya, Luzon Island (DMG, unpublished notes). These small ants share similar morphological features with Tetramorium, eg. a sharp clypeal ridge in front of antennal sockets and a lamella at the tip of the sting, but have a heart-shaped head, a broader, convex clypeus, and a keel-like petiolar venter. They are known to form large, ground-dwelling colonies nesting in rotten logs or in the ground. These ants may be collected by sifting leaf litter and pitfall trapping. Keys to species: Brown (1964) (World revision); Bolton (1976) (World tribal revision), (1986) (World revision).

Rhytidoponera

Genus Rhytidoponera Mayr, 1862 – Ectatomminae: 105 spp., at least 1 known from PH.

Brown (1958) reports one species from the Philippines, Rhytidoponera araneoides (Le Guillou, 1842). There are specimens of an unidentified species from “San Francisco, Agusan”, Mindanao Island, in the MCZ Ant Collection of Harvard University. These hard-bodied ants are superficially similar to Gnamptogenys but have a tooth at the anterolateral edge of the pronotum, and a different petiole shape. They are generalist predators or scavengers and nest in the ground under rocks but may also be arboreal. These ants may be collected by sifting leaf litter, pitfall trapping, beating low vegetation over a white sheet and inspecting tree hollows and dead branches in the canopy.

Romblonella

Genus Romblonella Wheeler, 1935a – Myrmicinae: 9 spp., 1 known from PH.

There is one valid species known from the Philippines, Romblonella opaca (F. Smith, 1861). It is a widespread species, having been collected in the far-apart islands of Romblon, Rapu-rapu, and Palawan (DMG, unpublished notes). These small ants have a large, bulbous, sessile petiole with a tiny denticle on the lower surface. They seem to be tolerant of human disturbance because the Rapu-rapu specimen was collected on a concrete sidewalk. They are ground-dwelling. These ants may be collected by sifting leaf litter, pitfall trapping, and hand collecting. Keys to species: Bolton (1976) (World revision), Taylor (1991a) (Australian).

Simopone

Genus Simopone Forel, 1891 – Cerapachyinae: 16 spp., 1 known from PH.

The unique holotype specimen of the only known species from the Philippines, Simopone chapmani Taylor, 1966, is deposited in the MCZ Ant Collection. These small ants bear a long barrel-shaped petiole and no tibial spurs on the middle legs. Simopone ants are often arboreal, and presumed to be predators on other ants. These ants may be collected by beating low vegetation over a white sheet and examining dead twigs or branches in the canopy.

Figure 26. Full-face and profile images of Philippine ant genera. Rhopalomastix species PH-01 queen (A, B). Rhoptromyrmex wroughtonii (C, D). Rhytidoponera species PH-01 (E, F).

Solenopsis

Genus Solenopsis Westwood, 1840 – Myrmicinae: 285 spp., at least 1 known from PH.

Ants of this genus exhibit 2 main lifeways: the fire ants live independently in familiar ant mounds while the tiny thief ants nest beside other ants or termites. The species known in the Philippines, Solenopsis geminata (Fabricius, 1804), was introduced by human commerce from the United States. S. geminata, also called fire ants, is a common pest in households and agricultural areas. They are dimorphic, with major and minor workers. There are unidentified specimens of Solenopsis subgenus Diplorhoptrum, which are tiny, monomorphic ants, collected by Perry Buenavente from a transect studies on Mt. Palali, Nueva Vizcaya Province, Luzon Island and 2 mountains in Mindanao Island (DMG, unpublished notes). There are also specimens of an unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). These ants may be collected by sifting leaf litter, pitfall trapping, and hand collecting. Key to species: Trager (1991) (World, geminata-group).

Stigmatomma

Genus Amblyopone Erichson, 1842 – Amblyoponinae: 69 spp., 3 known from PH.

These cryptic ants have characteristic elongate mandibles and a denticulate anterior clypeal border. In these ants, the gaster is broadly attached to the petiole. They are found under the bark of logs, in rotten wood, leaf litter, or soil and are known to prey on centipedes and beetle larvae. Key to species: Taylor (1979) (Melanesia). Genus Strumigenys F. Smith, 1860c – Myrmicinae: 474 spp., at least 20 known from PH.

There are specimens of an unidentified species from a transect study of Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). There are also specimens of unidentified species from Eastern Samar Province, Samar Island. These tiny to small, cryptic ants have opposing mandibles with teeth only at the apex, and which usually close with a visible gap. Similar to Pyramica, they also have spongy lobes at the sides of the propodeum, petiole, and postpetiole. They hunt for prey, usually Collembola (springtails), and nest in the leaf litter. These ants may be collected by inspecting dead that are stuck together for nests before sifting leaf litter. Key to species: Bolton (2000) (World revision).

Strumigenys

32 known from PH.

These small, slow-moving ants have opposing mandibles that have interlocking teeth and usually close without a gap. Similar to Strumigenys, they also have spongy lobes at the sides of the propodeum, petiole, and postpetiole. They hunt for prey and nest in the leaf litter. These ants may be collected by inspecting dead leaves stuck together before sifting the leaf litter and pitfall trapping. Key to species: Bolton (2000) (World revision).

Recurvidris

Genus Recurvidris Bolton, 1992 – Myrmicinae: 9 spp., at least 2 known from PH.

There are specimens of at least 2 unidentified species from a transect study in Mt. Isarog, Bicol Region, Luzon Island (Alpert and General in prep.). There are also specimens from Mindanao Island. Zettel (2008) recently described Recurvidris nigrans Zettel, 2008 from Negros Island. These small, slender ants have propodeal spines that curve upward and forward, and a ventrally expanded gaster. They are ground-dwelling and forage in the leaf litter. These ants may be collected by sifting leaf litter, underground baiting and pitfall trapping. Keys to species: Bolton (1992), Zettel (2008). Genus Tapinoma Foerster, 1850 – Dolichoderinae: 93 spp., 3 known from PH.

The species known from the Philippines include the widespread invasive species, T. melanocephalum, which commonly invades households in search of sweet food. These small ants have a small, flat petiole and reflexed apex of the gaster, such that only four segments are visible in dorsal view. They may forage in the foliage or on the ground for dead insects but also tend coccids and aphids for honeydew. They nest in just about any available cavity in soil, under stones and bark, in living plants, and houses. These ants may be collected by sifting leaf litter, beating low vegetation over a white sheet, and baiting with honey. Shattuck (1999), Fisher and Bolton (2007).

Figure 27. Full-face and profile images of Philippine ant genera. Romblonella opaca (A, B). Simopone conradti Emery, 1899 (N.B. Not from the Philippines, the unique holotype of S. chapmani is badly broken and cannot help the student visualize this rare genus) (C, D). Solenopsis geminata minor worker (E, F).

Technomyrmex

Genus Technomyrmex Mayr, 1872 – Dolichoderinae: 90 spp., 10 known from PH.

The species known from the Philippines include the widespread invasive species, T. albipes (F. Smith, 1861), which is very common in highly disturbed or agricultural areas. These ants also have a small, flat petiole but with five visible gastral segments in dorsal view. It can be difficult to distinguish them from Tapinoma, especially in shrunken or deformed specimens. They are scavengers and forage in the foliage and on the ground. They nest in the soil, in rotten wood, and under bark and rocks. These ants may be collected by sifting leaf litter, pitfall trapping, beating low vegetation over a white sheet, hand collecting, and flipping over rocks and loose bark. Key to species: Bolton (2007) (World revision).

Tetramorium

Genus Tetramorium Mayr, 1855 – Myrmicinae: 459 spp., at least 19 from PH.

The Philippine species include widespread invasive species such as T. pacificum Mayr, 1870 and T. tonganum Mayr, 1870, which are common in highly disturbed or agricultural areas. Schlick-Steiner et al. 2006 recently revived T. manobo (Calilung, 2000) from synonymy. There are specimens of several undescribed species (including a species misidentified as Leptothorax by Samson et al. 1997) from a transect study on Mt. Isarog, Bicol Region, Luzon Island (Alpert and General, in prep.). These small ants have a sharp clypeal ridge in front of the antennal sockets and a lamella at the tip of the sting, but the head is roughly rectangular in shape and with a straight-edged or slightly concave clypeus. They are variable in size, sculpture of the cuticle, and color. These ants are generalist predators and scavengers, foraging on the ground. They nest in twigs, rotten wood, under rocks or bark. These ants may be collected by sifting leaf litter, pitfall trapping, breaking into rotten wood and twigs, and flipping over rocks and loose bark. Keys to species: Bolton (1976, 1977) (Oriental, Indo-Australian), Schlick-Steiner et al. 2006.

Tetraponera

Genus Tetraponera F. Smith, 1852 – Pseudomyrmicinae: 118 spp., 9 known from PH.

These long and slender arboreal ants have large, somewhat flattened eyes, and a reduced clypeus, such that the antennal sockets are near the front edge of the head. They nest in dead twigs and branches. These ants may be collected by beating low vegetation over a white sheet and inspecting dead twigs and branches in the canopy. Key to species: Ward (2001) (Oriental, Australian).

Tyrannomyrmex

Genus Tyrannomyrmex Fernandez, 2003 – Myrmicinae: 3 sp., 1 known from PH.

This is a genus that is very rarely collected. The three known species were all described from unique specimens. A single male specimen, in the Philippine collection

Figure 28. Full-face and profile images of Philippine ant genera. Strumigenys chapmani Brown, 1954 (A, B). Tapinoma williamsi (Wheeler, 1935b) (C, D). Technomyrmex albipes (E, F).

of the Bernice P. Bishop Museum of Hawaii, appears to belong to this genus. If confirmed, it will be the first known male specimen of Tyrannomyrmex. These small ants have a curved petiole surmounted by a low node, and strongly-curved mandibles with only two teeth. They likely forage deep in the soil and only occasionally emerge in the leaf litter (Alpert 2011). These ants may be collected by underground baiting, sifting leaf litter and pitfall trapping. Fernandez (2003), Bolton et al. 2006, Alpert (2011).

Figure 29. Full-face and profile images of Philippine ant genera. Tetramorium khnum Bolton, 1977 (A, B). Tetraponera allaborans (Walker, 1859) (C, D). Tyrannomyrmex species PH-01 (E, F).

Vollenhovia

Genus Vollenhovia Mayr, 1865 –Myrmicinae: 72 spp., 6 known from PH.

These flat and slender ants have a large, keel-like ventral petiolar process. They nest in rotten wood and under rocks and forage in leaf litter. Little is known about their biology. These ants may be collected by sifting leaf litter, pitfall trapping, breaking into rotten wood and flipping over rocks. Bolton (2003).

Vombisidris

Genus Vombisidris Bolton, 1991 –Myrmicinae: 16 spp.,at least 2 known from PH.

A recently described species, Vombisidris philippina Zettel and Sorger, 2010, was collected from five locations on three different islands, representing the first record of this genus in the Philippines (Zettel and Sorger, 2010a). They also report at least another species, represented by a dealate queen from yet another location. These small ants are recognizable by a sinuate subocular groove running from the mandibular insertion to the laterooccipital margin of the head. Ants of this genus are arboreal, at least one species (V. humboldticola Zacharias and Rajan, 2004 from southern India) nesting in domatia, which are specialized swollen plant structures for harboring ants. These ants may be collected by beating low vegetation over a white sheet and inspecting domatia of epiphytes. Zettel and Sorger (2010a).

Unnamed genus PH-01

– Myrmicinae: 1 sp., 1 known from PH.

(New record). Specimens were extracted from berlesate from three locations on Samar Island. In Bolton (1994), this ant keys out to Mayriella, but bears little resemblance to it. The ant more closely resembles Tetheamyrma with its bidentate clypeal projection (S. Cover, pers. comm.), but has 10-segmented antennae and has no spatulate or lamellate hairs on the inner margin of the mandibles. Images are available online at: http://pick4.pick.uga.edu/mp/20q?act=x_antandpath=Insecta/Hymenoptera/Formicidae/Adelomyrmex/sp_phi1andname=Adelomyrmex+sp_phi1andxml=Ants_Philippinesandauthority=unknown+species. These ants may be collected by sifting leaf litter and pitfall trapping.

Unnamed genus PH-02

– Myrmicinae: 1 sp. (?), 1 known from PH.

(New record). There is a single specimen of this undescribed genus in the Philippine collection of the Bernice P. Bishop Museum in Honolulu, Hawaii. The collection was loaned to GDA, who found this unique specimen. The specimen was collected by L.W. Quate from San Francisco, Agusan del Sur in 1959. In Bolton (1994), this specimen keys out to “Undescribed genus” (p. 89) (P. Ward and S. Cover, pers. comm.). The diagnostic characteristic of this genus is the presence of the petiolar spiracle in the node, rather than in the peduncle. No collection information is included with the specimen, however, applying different collecting techniques may discover this species.

Figure 30. Full-face and profile images of Philippine ant genera. Vollenhovia ambitiosa Menozzi, 1925 (A, B). Vombisidris philippina (image reproduced with permission of NHMW and antbase.net) (C, D). Unnamed genus PH-01 (E, F).

Unnamed genus PH-03

– Myrmicinae: 2 spp., 2 known from PH.

(New record). A good nest series of this undescribed genus was collected by Perry Buenavente in a transect study of Mt. Palali, Nueva Vizcaya Province, Luzon Island. A unique specimen of another species (now lost due to theft, Taylor, pers. comm.) was earlier collected by Bambet Alto from Mt. Isarog. These reddish-orange ants seem to prefer high elevations, having been collected from >1450 masl at both locations. These ants may nest in rotten wood (DMG, unpublished notes). They are characterized by stemmed mandibles with blunt or rounded denticles and a subsessile petiole with an angle protecting the spiracle. These ants may be collected by breaking into rotten wood, sifting leaf litter and pitfall trapping.

Figure 31. Full-face and profile images of Philippine ant genera. Unnamed genus PH-02 (A, B). Unnamed genus PH-03 (C, D).

Key to Subfamily of Philippine Ants

Key to the subfamilies of Philippine ants, based on the worker caste (adapted from Bolton, 1994

Key to Amblyoponinae

Key to Philippine Amblyoponinae 1 Frontal lobes large, extending forward beyond anterior clypeal margin; antennal funiculus flattened or compressed Myopopone -Frontal lobes never extending beyond anterior clypeal margin; antennal funiculus more or less circular in cross-section 2

2(1) Mandibles with only 3 teeth, with median tooth smallest; at full closure, little or no gap between mandibles and anterior clypeal margin Prionopelta - Mandibles with more than 3 teeth; at full closure, large gap between mandibles and anterior clypeal margin 3

3(2) Posterior margin of head strongly concave; apex of mandible blunt or rounded Mystrium - Posterior margin of head at most weakly concave; apex of mandible acute Amblyopone

Key to Cerapachyinae

Key to Philippine Cerapachyinae

1 Antennal sockets set close together, separated by a narrow triangular posterior extension of the clypeus; mesotibial spurs present Cerapachys - Antennal sockets widely separated, set apart by a wide triangular posterior extension of the clypeus; mesotibial spurs absent Simopone

Key to Dolichoderinae

Key to Philippine Dolichoderinae

1 In profile, petiole overhung by anterior part of gaster and node absent 2 - In profile, petiole with a conspicuous node, and not overhung by gaster 3

2 (1) In profile, gaster with 4 dorsal segments (tergites), the fifth segment reflexed so that anal pore is ventral, not a termination of gaster Tapinoma - In profile, gaster with 5 dorsal segments (tergites) and anal pore is terminal Technomyrmex

3(1) Palps short and hard to see; in profile, dorsal face of propodeum much shorter than propodeal declivity, giving the mesosoma a compact appearance Bothriomyrmex - Palps long and conspicuous; dorsal face and declivity of propodeum about equal in length so that the mesosoma appears elongated 4

4(3) Head and mesosoma much longer than broad; legs extremely elongated Leptomyrmex - Head roughly triangular and mesosoma not elongated; legs not elongated 5

5(4) Mesosoma with thick and sculptured integument Dolichoderus - Mesosoma with thin integument and generally smooth or shagreened 6

6(5) In profile, rear face of propodeum concave; in side view, metanotal groove a narrow, distinct notch Ochetellus - In profile, rear face of propodeum convex, rarely flat; in side view, metanotal groove a broad impression 7

7(6) In frontal view, posterior margin of head strongly concave and eyes located anterior to midline of of head; petiolar node inclined forward; polymorphic workers Philidris - In frontal view, posterior margin of head at most weakly concave, usually flat or convex; eyes posterior to midline of head; petiolar node more or less vertical; monomorphic workers Iridomyrmex

Key to Etatomminae

Key to Philippine Ectatomminae

1 In profile, anteroventral margin of pronotum with distinct tooth; hind pretarsal claw with median tooth Rhytidoponera - In profile, anteroventral margin of pronotum rounded or bluntly angulate; hind pretarsal claw without median tooth Gnamptogenys

Key to Formicinae

Key to Philippine Formicinae

1 Antenna 8-segmented, folding back below eye; eyes always large Gesomyrmex - Antenna with 9- to 12-segmented, folding back above eye; eyes variable in size 2

2 (1) Antenna 9- to 11-segmented 3 - Antenna 12-segmented 6

3 (2) Propodeum and petiole armed with a pair of spines or teeth Lepisiota - Propodeum and petiole without spines or teeth 4

4(3) Palps very short and extremely difficult to see; eyes minute Acropyga - Palps long and prominent; eyes usually well-developed, often large 5

5 (4) Antennal scapes extremely long, extending more than half their length beyond posterior margin of head; erect hairs absent on dorsum of mesosoma Anoplolepis - Antennal scape, notably shorter, seldom extending much beyond posterior margin of head; erect hairs sometimes present on dorsum of mesosoma Plagiolepis

6 (2) Mandibles with 10 or more teeth or denticles 7 - Mandibles with fewer than 10 teeth or denticles 8

7(6) Mandibles linear and longer than head length, with sharp teeth along most of their length and crossing at apices when closed; eyes enormous Myrmoteras - Mandibles triangular and shorter than head length; eyes large but not taking up most of the sides of head Oecophylla

8(6) Antennal sockets almost abutting posterior clypeal margin; ring of hairs present around acidopore 9 - Antennal sockets well posterior to posterior clypeal margin; ring of hairs often absent around acidopore 15

9(8) Maxillary palp short and inconspicuous, with 2–4 segments 10 - Maxillary palp long and conspicuous, with 6 segments 11

10(9) In side view, mesonotal constriction present; mandibles strongly curved Euprenolepis - In side view, mesonotal constriction absent; mandibles not strongly curved Pseudolasius

11(9) Mesosoma and head without coarse erect hairs Overbecia - Mesosoma and head with coarse erect hairs 12

12(11) In side view, mesosoma long and slender, with or without constriction of mesonotum 13 - In side view, mesosoma short and compact, without constriction of mesonotum 14

13(12) In side view, pronotum only slightly convex; erect setae on head randomly scattered on surface Paratrechina - In side view, pronotum convex; erect setae on head form 2 parallel rows Prenolepis

14(12) Propodeum with 1 pair of erect setae Paraparatrechina - Propodeum without a pair of erect setae Nylanderia

15(8) In side view, metathoracic spiracles forming turbercles that are the highest prominences of the mesosoma Forelophilus - In side view, metathoracic spiracles not forming tubercles that are the highest prominences of the mesosoma 16

16(15) Petiole node lacking teeth or spines; first gastral tergite distinctly less than half total length of gaster Camponotus - Petiole node armed with spines, teeth, or denticles; first gastral tergite large, at least half of total length of gaster 17

17(16) Spines or teeth usually present on pronotum, propodeum, or both; body usually covered with short appressed hairs and some erect hairs Polyrhachis - Spines or teeth absent from pronotum and propodeum, often present on petiole; body usually densely covered with long erect hairs Echinopla

Key to Myrmicinae

Key to Philippine Myrmicinae

1 In side view, antennal scrobe present below eye 2 - In side view, antennal scrobe absent or present above eye 3

2 (1) Antenna 7-segmented; antennal scape triangular, widest near insertion and tapering distally; petiole pedunculate; propodeum armed with thin longitudinal flanges Eurhopalothrix - Antenna 11-segmented; antennal scape rod-like; petiole sessile; propodeum armed with 2 thick spines Cataulacus

3 (1) Petiole lacking a distinct node 4 - Petiole with a distinct node 6

4 (3) Petiole transversely flattened; in dorsal view, gaster roughly heart-shaped; petiole attached to dorsal surface of gaster Crematogaster - Petiole roughly barrel-shaped; in dorsal view, gaster not heart-shaped 5

5 (4) Side of head with a longitudinal ridge running below the eye, from rear border to mandibular insertion; lobes of posterior margin of the head not produced into blunt points Myrmecina - Side of head lacking a ridge below the eye; lobes of the posterior margin of the head produced into blunt points Dilbocondyla

6 (3) Mandibles bidentate; mesosoma without a distinct transverse suture Tyrannomyrmex - Mandibles with more than 2 teeth; mesosoma with a distinct transverse suture 7

7 (6) Petiole and postpetiole with at least some light-colored sponge-like tissue 8 - Petiole and postpetiole never with sponge-like tissue 10

8(7) Sponge-like tissue only on ventral surface of petiole, postpetiole, and the 4th abdominal segment; antenna 11-segmented Dacetinops - Sponge-like tissue on lateral, as well as ventral, surfaces of petiole, postpetiole, and 4th abdominal segment; antenna 4- to 6-segmented 9

9(8) Mandibles usually linear, at full closure, separated by a gap and with teeth only at distal end of mandibles Strumigenys - Mandibles triangular, at full closure, slightly overlapping and with teeth along entire inner margin Pyramica

10 (7) Antenna with a distal club of 2 segments 11 - Antenna with a distal club of 3 or 4 segments, or with an indistinct club 16

11 (10) Antennal scrobe present; head and mesosoma punctate Mayriella - Antennal scrobe absent; head and mesosoma smooth, rugose, or reticulate 12

12 (11) Head and mesosoma sculptured; petiole at most with a short peduncle 13 - Head and mesosoma smooth between sparse punctures; petiole pedunculate 14

13 (12) Head and mesosoma longitudinally rugose; propodeum unarmed Rhopalomastix - Head and mesosoma strongly reticulate; propodeum bidentate Unnamed genus PH-01

14 (12) Propodeum smooth and unarmed; anterior clypeal margin with two angles, or teeth, bearing hairs Solenopsis - Propodeum with a pair of spines or teeth; anterior clypeal margin simple; workers dimorphic or polymorphic 15

15 (14) Clypeus with 2 longitudinal carinae immediately anterior to antennal sockets; workers dimorphic Carebara - Clypeus smooth and without longitudinal carinae; workers polymorphic Pheidologeton

16(10) Antenna with 7–9 segments 17 - Antenna with 10–12 segments 18

17 (16) Antenna 7-segmented; mesosoma strongly reticulate; propodeal spines long; anterior peduncle of petiole as long as petiolar node height Myrmicaria - Antenna 9-segmented; mesosoma weakly reticulate and shield-like; propodeal spines short; anterior peduncle shorter than petiolar node height Meranoplus

18 (16) Eyes absent 19 - Eyes present, even if only one ommatidium 20

19(18) Petiole and postpetiole with prominent anteroventral processes; integument firm and sclerotized Liomyrmex - Petiole and postpetiole lacking anteroventral processes; integument fragile and poorly sclerotized Anillomyrma

20 (18) In dorsal view, postpetiole swollen and at least twice as wide as petiole; antenna 12-segmented Cardiocondyla - In dorsal view, postpetiole as wide as, or slightly wider than, petiole; antenna 10- to 12-segmented 21

21 (20) Clypeus with an anterior forked extension overhanging the mandibles; body covered with evenly-spaced clavate hairs Calyptomyrmex - Clypeus without an anterior forked extension; body hairs not clavate 22

22 (21) Sides of clypeus immediately anterior to antennal sockets produced into a sharp ridge 23 - Sides of clypeus immediately anterior to antennal sockets not produced into a sharp ridge 24

23 (22) In frontal view, head roughly heart-shaped, with posterior margin of the head strongly concave; antenna 12-segmented Rhoptromyrmex - In frontal view, head roughly rectangular, with posterior margin of the head at most weakly concave; antenna 10- to 12-segmented Tetramorium

24 (22) Anterior clypeal margin with a single median seta extending over the mandibles; propodeum at most with blunt angles; antenna 10- to 12-segmented Monomorium - Anterior clypeal margin with a pair of setae, or a series of long, strong setae, or hairless 25

25 (24) Antenna 11-segmented 26 - Antenna 12-segmented 31

26 (25) Antennal sockets completely exposed; anterior clypeal margin denticulate Pristomyrmex - Antennal sockets partially or completely covered by frontal lobes; anterior clypeal margin usually simple 27

27 (26) Antennal scrobes deep and narrow, overhung by expanded frontal carinae Mtapone - Antennal scrobes absent or very shallow 28

28 (27) Propodeum unarmed; in profile, petiole lacking peduncle and usually with a large subpetiolar process Vollenhovia (part) - Propodeum spinose 29

29 (28) Propodeal spines curving upward and forward Recurvidris - Propodeal spines straight 30

30(29) Propodeal spines long and sharp; in side view, pronotum convex, much higher than propodeum Lophomyrmex - Propodeal spines short and blunt; in side view, pronotum flat, almost level with propodeum Gauromyrmex

31 (25) Antenna, gently expanding distally, with a weak to indistinct 4-segmented club Aphaenogaster - Antenna abruptly expanding distally, with a distinct 3-segmented club 32

32(31) Petiole lacking peduncle 33 - Petiole pedunculate 35

33 (32) Propodeum armed with stout spines (curved in dorsal view); petiole armed with a denticle ventrally Romblonella - Propodeum unarmed or at most with denticles 34

34 (33) Petiole with a conspicuous ventral process; mandibles curved mesad and without a stem at insertion Vollenhovia (part) - Petiole without a conspicuous ventral process; mandibles linear-triangular and with a stem at insertion (see diagnosis in “Brief Generic Accounts”) Unnamed genus PH-03

35 (33) Head enormous, obscuring pronotum in dorsal view; anterior clypeal margin overhanging mandibles and armed with blunt denticles or a median rectangular extension Acanthomyrmex - Head does not obscure pronotum in dorsal view; anterior clypeal margin simple and not overhanging mandibles 36

36 (35) In profile, mesosomal dorsum with a flat or weakly convex outline; petiolar node blocky or roughly rectangular Paratopula - In profile, dorsal mesosomal dorsum with a strongly convex outline; petiolar node globular or rounded 37

37(36) In profile, petiolar spiracle located on node, not on peduncle (see diagnosis in “Brief Generic Accounts”) Unnamed genus PH-02 - In profile, petiolar spiracle located on peduncle 38

38 (37) Mandible with 5 teeth with a long diastema between 3rd and 4th tooth; side of head usually with a sinuate longitudinal groove running from the posterior margin to the mandibular insertion Vombisidris - Mandibular dentition not as above; sinuate longitudinal groove at side of head absent 39

39 (38)In profile, petiole with a short anterior peduncle; mandibles dentate; workers monomorphic Lordomyrma - In profile, petiole with a long anterior peduncle; mandibles dentate or toothless; workers dimorphic Pheidole

Key to Ponerinae

Key to Philippine Ponerinae

1 Mandibles long and linear, inserted at middle of the anterior margin of head 2 - Mandibles triangular or linear, inserted at sides of head 3

2 (1) Frons with a conspicuous groove running along the midline; in profile, petiole node produced into a dorsal sharp point Odontomachus - Frons simple, at most with shallow striations; in profile, petiole node blunt Anochetus

3 (1) Antennal insertions well separated; hind tibia with two pectinate spurs Platythyrea - Antennal insertions closely approximated; hind tibia with one pectinate spur, but may have a smaller simple spur in front of it 4

4 (3) Hind tibia with only one pectinate spur, without a smaller, simple anterior spur 5 - Hind tibia with two spurs with the smaller spur simple 8

5 (4) Side of mandible near insertion with a small circular pit Pachycondyla (part) - Side of mandible near insertion without a pit 6

6 (5) Outer surface of middle tibia and middle and hind tarsi with strong, peg-like teeth; pronotum flat and shelf-like in profile Centromyrmex - Outer surface of middle tibia and middle and hind tarsi with hairs, but never with strong, peg-like teeth; pronotum convex in profile 7

7(6) Subpetiolar process with an oval or circular translucent window, and with a sharp posterior angle Ponera - Subpetiolar process without an oval or circular translucent window, and usually blunt or rounded posteriorly Hypoponera

8 (4) Tarsal claws on hind leg either pectinate or with one or more teeth on inner surface 9 - Tarsal claws on hind leg simple, never with teeth on inner surface 10

9 (8) Ocelli present; mandibles long and forceps-like with a triangular flange beneath Harpegnathos - Ocelli absent; mandibles variable but never long and forceps-like with a triangular flange beneath Leptogenys

10 (8) Petiole node with a pair of spines Diacamma - Petiole node simple 11

11 (10) Side of pronotum with a pair of large blunt angles; anterior clypeal margin with small blunt teeth or denticles Odontoponera - Side of pronotum and anterior clypeal margin simple 12

12 (11) Mandibles, when fully closed, with a large gap between them; eyes situated very near base of mandibles Myopias - Mandibles, when fully closed, slightly overlap along the inner margin; eyes situated away from base of mandibles Pachycondyla (part)

Key to Proceratiinae

Key to Philippine Proceratiinae

1 Fourth abdominal segment straight or slightly curved, so that apex of gaster is directed posteriorly Probolomyrmex - Fourth abdominal segment strongly curved, so that apex of gaster is directed anteriorly 2

2 (1) Apical segment of antenna extremely large and bulbous; antennal sockets on shelf protruding over mandibles Discothyrea - Apical segment moderately enlarged, but not bulbous; antennal sockets not protruding over mandibles Proceratium

Glossary

• Abdominal segments – morphologically, segment I of the ant abdomen is the propodeum (see below), followed by the petiole (see below), which is segment II. The postpetiole (see below), when present, is segment III. Otherwise, the first segment of the gaster (see below) is segment III.
• Acidopore – at the apex of the gaster (see below), a round, somewhat raised, orifice, often fringed with hairs at the tip; diagnostic of ants belonging to subfamily Formicinae.
• Angle – a triangular, broad-based, tooth-like extension of the cuticle.
• Antennal club – describes an antennal funiculus that enlarges apically, to form a distinct club of 2-4 segments.
• Antennal funiculus – the portion of the antenna distal to the antennal scape; composed of 3-12 segments of varying size.
• Antennal scape – the elongated basal segment of the antenna.
• Antennal scrobe – a groove above or below the eye, which accepts the folded antenna.
• Anterior clypeal margin – the leading edge of the clypeus (see below).
• Bidentate – armed with 2 teeth (mandibles) or short spines (propodeum).
• Clavate – refers to hairs that are blunt and club-shaped.
• Clypeus – the anterior sclerite (see below) of the dorsal head, which consists of narrow lateral portions and a shield-like median portion.
• Coxa – see Leg segments.
• Declivity – downward slope, eg. as of the propodeum.
• Dentate – describes mandibles armed with teeth.
• Denticle – a small triangular tooth, much reduced in size.
• Denticulate – armed with denticles.
• Edentate – describes mandibles without teeth or denticles on the inner (mastigatory) margin.
• Epigaeic – describes ants that live above ground (compare with hypogaeic).
• Filiform – describes an antennal funiculus with segments of approximately the same size (compare with antennal club).
• Frons – the region of the head extending from behind the clypeus, and between the eyes to the posterior margin of the head.
• Frontal carina – (pl. carinae) thin ridges of cuticle on the front of the head, which may form the dorsal margins of the antennal scrobes (see above).
• Frontal lobe – projection of the frontal carina (see above), which may partially or completely cover the antennal sockets.
• Gaster – old name for the third main body division of the ant body.
• Head – the first main body division of the ant body.
• Hypogaeic – describes ants that live underground (compare with epigaeic).
• Labial palps – the segmented sensory appendages attached to the labium, found on the anteroventral surface of the head, with a maximum of four segments. These are the inner pair of palps.
• Leg segments – Each leg consists of the basal coxa, a small trochanter, a usually long and swollen femur, a tibia, and a 5-segmented tarsus, ending in a pair of tarsal claws. Pro-, meso-, and meta- are prefixes that indicate the leg segment of the particular thoracic segment.
• Mandibles – anterior appendages of the head, with which the ant manipulates its environment. They are variable in shape, dentition, and function, and extremely important in ant taxonomy.
• Maxillary palps – the segmented sensory appendages attached to the maxillae, found on the anteroventral surface of the head. With a maximum of six segments, they are usually the longer and larger of the palps and are the outer pair of palps.
• Mesosoma – (= alitrunk) the second main body division of the ant body. Morphologically, it is composed of the three thoracic segments (pro-, meso-, and metathorax) to which is fused the propodeum (see below).
• Mesonotum – the second tergite (see below) of the mesosoma (see above).
• Mesotibia – see Leg segments.
• Metanotal groove – a transverse depression between the mesonotum (see above) and the propodeum (see below).
• Metapleural gland – an exocrine gland found on the posteroventral side of the mesosoma (see above), just above the metacoxa (see Leg segments). This is one of the defining features of ants, although it has been secondarily lost or reduced in some taxa.
• Node – a prominent bulge on the dorsal surface of the petiole.
• Ocellus – (pl. ocelli) a simple eye found on the frons (see above) of the head, usually in reproductives but may also be found in the workers of some taxa.
• Ommatidium – an individual facet of the compound eye.
• Palp formula – a standardized method of giving the number of segments of the maxillary and labial palps. The number of maxillary palp (see above) segments is given first, followed by the number of labial palp (see above) segments; thus “PF 6,4” indicates that there are six maxillary palp segments and four labial palp segments.
• Pectinate – describes tarsal claws or tarsal spurs that are comb-like.
• Pedunculate – bearing a stem-like projection, or peduncle, that is anterior to the petiolar node (compare with sessile and sub-sessile).
• Petiole – the isolated segment separating the mesosoma (see above) and the gaster (see above). This is one of the defining features of ants. Morphologically, it is the second segment of the abdomen.
• Postpetiole – the second isolated and reduced segment separating the mesosoma (see above) and the gaster (see above), in ants with a 2-segmented waist. Morphologically, it is the third abdominal segment.
• Pronotum – the first tergite (see below) of the mesosoma (see above).
• Propodeum – (= epinotum) the dorsal posterior plate of the mesosoma (see above). Morphologically, it is the first segment of the abdomen, fused to the thorax. It may have specializations such as spines, teeth, or lobes.
• Punctate – describes surface sculpturing composed of round pits which may be shallow or deep.
• Reticulate – describes surface sculpturing composed of an irregular network of thin ridges of cuticle.
• Rugose – describes surface sculpturing of raised ridges without cross-ridges.
• Sclerite – a hardened or stiffened plate of the integument.
• Sculpturing – features on the body surface which may be pits, grooves, striae, ridges, punctures, or a combination.
• Sessile – describes a petiole without a peduncle, such that the petiolar node is very close to the propodeum (compare with pedunculate).
• Shagreened – describes body surface that has a dull, light-absorbing quality.
• Spinose – armed with spines.
• Sternite – the lower or ventral sclerite (see above) of a segment.
• Striate – describes sculpturing composed of shallow, parallel grooves or lines.
• Tergite – the upper or dorsal sclerite (see above) of a segment.

Acknowledgments

We would like to thank Fr. Joel E. Tabora, S.J., president of Ateneo de Naga University, for lending the ant specimens to the Museum of Comparative Zoology on long-term basis, and Emelina G. Regis (INECAR), for allowing us the use of her office space and laboratory equipment during our transect work. We also thank Joanaviva Caceres-Plopenio (Ateneo de Naga University), Perry Buenavente (Ateneo de Manila University), Arvin Diesmos and Ven Samarita (Philippine National Museum, Manila), Ireneo Lit, Jr. and Leonila A. C. Raros (UPLB-Museum of Natural History), Herbert Zettel (Museum of Natural History, Vienna), Ted Schultz (Smithsonian Institution), Hendrik Freitag, and Keith Arakaki (Bernice P. Bishop Museum) for giving us access to their specimens. For permitting us to use their images, we thank Katsuyuki Eguchi for Anillomyrma decamera, first published in Myrmecological News (Eguchi et al. 2010), Hans Peter Ravn for Noonilla copiosa, first published in Entomologiske Meddelelser (Petersen 1968), Martin Pfeiffer, NHMW and www.antbase.net for Vombisidris philippina, first published in Entomologica Austriaca (Zettel and Sorger 2010a). We are also very grateful to Denise R. General for post-processing the images and preparing the image plates. We also thank Stefan Cover and the late Roy Snelling for reviewing our paper.

References

This bibliography includes the reference to each original description of genus and species of Philippine ants listed in the Appendix as well as relevant publications of wider scope. Letters after the publication year simply refer to multiple publications in the same year.

• Agosti, D. 1992. Revision of the ant genus Myrmoteras of the Malay Archipelago (Hymenoptera, Formicidae). Rev. Suisse Zool. 99: 405-429.

• Agosti, D.; Majer, J. D.; Alonso, L. E.; Schultz, T. R. (eds.) 2000. Ants. Standard methods for measuring and monitoring biodiversity. Washington: Smithsonian Institution Press, xix + 280 pp.

• Alpert, G. D. 2007. A review of the ant genus Metapone Forel from Madagascar, pp. 8-18. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80.

• Alpert GD, General DM (2008) Philippine Ants. Antweb. Available at http://www.antweb.org/philippines.jsp [accessed 14.I.2011].

• Alpert GD, Cover SP, General DM, Samarita V (2006) Ants of the Philippines. Polistes Foundation, Discover Life. Available at http://pick4.pick.uga.edu/mp/20q?act=x_checklistandguide=Ants_Philippines [accessed 14.I.2011].

• André, E. 1892a. Matériaux myrmécologiques. Rev. Entomol. (Caen) 11: 45-56.

• Ashmead, W. H. 1904a. A list of the Hymenoptera of the Philippine Islands, with descriptions of new species. J. N. Y. Entomol. Soc. 12:1-22.

• Ashmead WH (1904b) Descriptions of new genera and species of Hymenoptera from the Philippine Islands. Proceedings of the US Philippine National Museum 28: 127–158.

• Ashmead, W. H. 1905b. New Hymenoptera from the Philippines. Proc. U. S. Natl. Mus. 29: 107-119.

• Baltazar, C. R. 1966. A catalogue of Philippine Hymenoptera (with a bibliography, 1758-1963). Pac. Insects Monogr. 8: 1-488.

• Baroni Urbani, C. 1971b. Einige Homonymien in der Familie Formicidae (Hymenoptera). Mitt. Schweiz. Entomol. Ges. 44: 360-362.

• Baroni Urbani, C. 1975a. Primi reperti del genere Calyptomyrmex Emery nel subcontinente Indiano. Entomol. Basil. 1: 395-411 PDF'

• Baroni Urbani, C. 1977a. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basel. Hymenoptera: Fam. Formicidae Genus Mayriella. Entomol. Basil. 2: 411-414.

• Baroni Urbani, C. 1977c. Materiali per una revisione della sottofamiglia Leptanillinae Emery (Hymenoptera: Formicidae). Entomol. Basil. 2: 427-488.

• Baroni Urbani, C.; De Andrade, M. L. 2003a. The ant genus Proceratium in the extant and fossil record (Hymenoptera: Formicidae). Mus. Reg. Sci. Nat. Monogr. (Turin) 36: 1-492

• Bernard, F. 1953b [1952]. La réserve naturelle intégrale du Mt Nimba. XI. Hyménoptères Formicidae. Mém. Inst. Fr. Afr. Noire 19: 165-270.

• Bihn, J.H. & Verhaagh, M. 2007. A review of the genus Mystrium in the Indo-Australian region. Zootaxa 1642: 1-12.

• Bingham, C. T. 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. London: Taylor and Francis, 506 pp.

• Bolton, B. 1974a. A revision of the Palaeotropical arboreal ant genus Cataulacus F. Smith (Hymenoptera: Formicidae). Bull. Br. Mus. (Nat. Hist.) Entomol. 30: 1-105.

• Bolton, B. 1975c. The sexspinosa-group of the ant genus Polyrhachis F. Smith (Hym. Formicidae). J. Entomol. Ser. B 44: 1-14.

• Bolton, B. 1976. The ant tribe Tetramoriini (Hymenoptera: Formicidae). Constituent genera, review of smaller genera and revision of Triglyphothrix Forel. Bull. Br. Mus. (Nat. Hist.) Entomol. 34: 281-379.

• Bolton, B. 1977. The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Oriental and Indo-Australian regions, and in Australia. Bull. Br. Mus. (Nat. Hist.) Entomol. 36: 67-151.

• Bolton, B. 1981a. A revision of the ant genera Meranoplus F. Smith, Dicroaspis Emery and Calyptomyrmex Emery (Hymenoptera: Formicidae) in the Ethiopian zoogeographical region. Bull. Br. Mus. (Nat. Hist.) Entomol. 42: 43-81.

• Bolton, B. 1986a. A taxonomic and biological review of the tetramoriine ant genus Rhoptromyrmex (Hymenoptera: Formicidae). Syst. Entomol. 11: 1-17.

• Bolton, B. 1988b. A review of Paratopula Wheeler, a forgotten genus of myrmicine ants (Hym., Formicidae). Entomol. Mon. Mag. 124: 125-143.

• Bolton, B. 1991. New myrmicine genera from the Oriental Region (Hymenoptera: Formicidae). Syst. Entomol. 16: 1-13.

• Bolton, B. 1992b. A review of the ant genus Recurvidris (Hym.: Formicidae), a new name for Trigonogaster Forel. Psyche (Camb.) 99: 35-48.

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

• Bolton B (2000) The ant tribe Dacetini. Memoirs of the American Entomological Institute 65: 1–1028.

• Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp.

• Bolton, B. 2007b. Taxonomy of the dolichoderine ant genus Technomyrmex Mayr (Hymenoptera: Formicidae) based on the worker caste. Contributions of the American Entomological Institute 35(1): 1-149.

• Bolton B, Alpert GD, Ward PS, Naskrecki P (2006) Bolton’s catalogue of the ants of the world: 1758–2005. Harvard University Press. (CD-ROM).

• Brown, W. L., Jr. 1948b. Strumigenys karawajewi, new name for a Sumatran ant. Entomol. News 59: 44.

• Brown, W. L., Jr. 1953g. Revisionary studies in the ant tribe Dacetini. Am. Midl. Nat. 50: 1-137.

• Brown, W. L., Jr. 1954j. The Indo-Australian species of the ant genus Strumigenys Fr. Smith: S. chapmani new species. Psyche (Camb.) 61: 68-73.

• Brown, W. L., Jr. 1957h ("1956"). The Indo-Australian species of the ant genus Strumigenys Fr. Smith: Three new Philippine species. Psyche (Cambridge) 63:113-118. [1957-11-18]

• Brown, W. L., Jr. 1958g. Contributions toward a reclassification of the Formicidae. II. Tribe Ectatommini (Hymenoptera). Bulletin of the Museum of Comparative Zoology 118: 173-362.

• Brown, W. L., Jr. 1960a. Contributions toward a reclassification of the Formicidae. III. Tribe Amblyoponini (Hymenoptera). Bulletin of the Museum of Comparative Zoology 122: 143-230.

• Brown, W. L., Jr. 1964e. Genus Rhoptromyrmex, revision of, and key to species. Pilot Regist. Zool. Card No. 11.

• Brown, W. L., Jr. 1973b. A comparison of the Hylean and Congo-West African rain forest ant faunas. Pp. 161-185 in: Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Washington, D.C.: Smithsonian Institution Press, viii + 350 pp.

• 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: 1-115

• Brown, W. L., Jr. 1976c. Contributions toward a reclassification of the Formicidae. Part VI. Ponerinae, tribe Ponerini, subtribe Odontomachiti. Section A. Introduction, subtribal characters. Genus Odontomachus. Stud. Entomol. 19: 67-171.

• Brown WL Jr (1978) Contributions toward a reclassification of the Formicidae. Part 6. Ponerinae, tribe Ponerini, subtribe Odontomachiti. Section B. Genus Anochetus and bibliography. Studia Entomologica (NS) 20: 549–652.

• Brown WL Jr (2000) [Untitled. Mitis–group. Pyramica mitis Brown sp. n.] 441–443 in: Bolton B. The ant tribe Dacetini. Memoirs of the American Entomological Institute 65: 1–1028.

• Brown, W. L., Jr.; Kempf, W. W. 1960. A world revision of the ant tribe Basicerotini. Stud. Entomol. (n.s.) 3: 161-250.

• Brown, W. L., Jr.; Wilson, E. O. 1957a. Dacetinops, a new ant genus from New Guinea. Breviora 77: 1-7.

• Calilung, M.V.J. 2000. A new genus, two new species and a new subspecies of Philippine ants. Philipp. Entomol. 14: 65-73.

• Cameron P (1886) On a new species of Strumigenys (S. lewisi) from Japan. Proceedings of the Manchester Literary and Philosophical Society 25: 229–232.

• Chapman, J. W. 1963. Some new and interesting Philippine ants (Hymenoptera: Formicidae). Philipp. J. Sci. 92: 247-263.

• Chapman, J. W.; Capco, S. R. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia. Monogr. Inst. Sci. Technol. Manila 1: 1-327.

• Chapman JW, Chapman E (1947) Escape to the hills. Jaques Cattell Press, Lancaster, PA, USA, 247 pp.

• Cole, A. C., Jr. 1949a. A study of the genus Gesomyrmex Mayr, and a description of a species new to the genus (Hymenoptera: Formicidae). Ann. Entomol. Soc. Am. 42: 71-76.

• Creighton, W. S. 1929a. New forms of Odontoponera transversa. Psyche (Camb.) 36: 150-154.

• Curtis, J. 1829. Myrmecina Latreillii. Plate 265 [plus 2 unnumbered pages of text] in: Curtis, J. British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland. Volume 6. London: published by the author, plates 242-289.

• Donisthorpe, H. 1937b. A new species of Harpegnathos Jerd., with some remarks on the genus, and other known species (Hym. Formicidae). Entomol. Mon. Mag. 73: 196-201.

• Donisthorpe, H. 1938d. New species and varieties of ants from New Guinea. Ann. Mag. Nat. Hist. 11(1): 593-599.

• Donisthorpe, H. 1941a. Lordomyrma niger sp. n. (Hym., Formicidae), with a key and notes on the genus. Entomol. Mon. Mag. 77: 36-38.

• Donisthorpe, H. 1941c. Descriptions of new species of ants from New Guinea. Ann. Mag. Nat. Hist. 11(7): 129-144.

• Donisthorpe, H. 1941. Descriptions of new ants (Hym., Formicidae) from various localities. Annals and Magazine of Natural History (11)8:199-210.

• Donisthorpe, H. 1942b. Descriptions of a few ants from the Philippine Islands, and a male of Polyrhachis bihamata Drury from India. Ann. Mag. Nat. Hist. 11(9): 64-72.

• Donisthorpe, H. 1943a. Descriptions of new ants, chiefly from Waigeu Island, N. Dutch New Guinea. Ann. Mag. Nat. Hist. 11(10): 167-176.

• Donisthorpe, H. 1943f. Myrmecological gleanings. Proc. R. Entomol. Soc. Lond. Ser. B 12: 115-116.

• Donisthorpe, H. 1947e. Some new ants from New Guinea. Ann. Mag. Nat. Hist. 11(14): 183-197.

• Dorow, W. H. O. 1995. Revision of the ant genus Polyrhachis Smith, 1857 (Hymenoptera: Formicidae: Formicinae) on subgenus level with keys, checklist of species and bibliography. Cour. Forschungsinst. Senckenb. 185: 1-113.

• Dorow, W. H. O.; Kohout, R. J. 1995. A review of the subgenus Hemioptica Roger of the genus Polyrhachis Fr. Smith with description of a new species (Hymenoptera: Formicidae: Formicinae). Zool. Meded. (Leiden) 69: 93-104.

• Dorow, W.H.O. & Maschwitz, U.& S. Rapp. 1990. The natural history of Polyrhachis (Myrmhopla) muelleri Forel 1893 (Formicidae: Formicinae), a weaver ant with mimetic larvae and an unusual nesting behavior. Tropical Zoology 3: 181-190.

• Drury D (1773) Illustrations of natural history, wherein are exhibited upwards of two hundred and twenty figures of exotic insects, according to their different genera. Vol 2B. White, London, vii + 90 pp.

• Eguchi, K. 2001b. A taxonomic study on Asian Pheidole (Hymenoptera, Formicidae): new synonymy, rank changes, lectotype designations and redescriptions. Insecta Koreana 18: 1-35.

• Eguchi, K. 2001a. A revision of the Bornean species of the ant genus Pheidole (Insecta: Hymenoptera: Formicidae: Myrmicinae). Tropics Monogr. Ser. 2: 1-154.

• Eguchi, K. 2003. A Study on the Male Genitalia of Some Asian Species of Pheidole (Hymenoptera, Formicidae, Myrmicinae). Sociobiology 41(2): 317-355PDF

• Eguchi, K. 2004. Taxonomic revision of two wide-ranging Asian ants, Pheidole fervens and P. indica (Insecta: Hymenoptera, Formicidae), and related species. Ann. Naturhist. Mus. Wien B. Bot. Zool. 105(B): 189-209.

• Eguchi, K., T. V. Bui, D. M. General and G. D. Alpert. 2009. Revision of the ant genus Anillomyrma Emery, 1913 (Hymenoptera: Formicidae: Myrmicinae: Solenopsidini). Myrmecological News 13: 31-36.

• Eguchi, K., Yoshimura, M. & Yamane, S. 2006. The Oriental species of the ant genus Probolomyrmex. Zootaxa 1376: 1-35.

• Emery, C. 1869b. Enumerazione dei formicidi che rinvengonsi nei contorni di Napoli con descrizioni di specie nuove o meno conosciute. Ann. Accad. Aspir. Nat. Secunda Era 2: 1-26.

• Emery, C. 1869c. Descrizione di una nuova formica italiana. Annu. Mus. Zool. R. Univ. Napoli 5: 117-118PDF

• Emery, C. 1870. Studi mirmecologici. Bull. Soc. Entomol. Ital. 2: 193-201.

• Emery, C. 1884a. Materiali per lo studio della fauna Tunisina raccolti da G. e L. Doria. III. Rassegna delle formiche della Tunisia. [part]. Ann. Mus. Civ. Stor. Nat. 21[=(2)(1): 373-384.

• Emery, C. 1887a [1886]. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia. [part]. Ann. Mus. Civ. Stor. Nat. 24[=(2)(4): 209-240.

• Emery, C. 1887b [1886]. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia. [part]. Ann. Mus. Civ. Stor. Nat. 24[=(2)(4): 241-258.

• Emery, C. 1887e. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia (continuazione e fine). [part]. Ann. Mus. Civ. Stor. Nat. 25[=(2)(5): 427-432.

• Emery, C. 1887f. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia (continuazione e fine). [part]. Ann. Mus. Civ. Stor. Nat. 25[=(2)(5): 433-448.

• Emery, C. 1887g. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia (continuazione e fine). [part]. Ann. Mus. Civ. Stor. Nat. 25[=(2)(5): 449-464.

• Emery, C. 1887h. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia (continuazione e fine). [concl.]. Ann. Mus. Civ. Stor. Nat. 25[=(2)(5): 465-473.

• Emery, C. 1889c. Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885-87). [part]. Ann. Mus. Civ. Stor. Nat. 27[=(2)(7): 485-512.

• Emery, C. 1889d. Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885-87). [concl.]. Ann. Mus. Civ. Stor. Nat. 27[=(2)(7): 513-520.

• Emery, C. 1890c. Studii sulle formiche della fauna neotropica. Bull. Soc. Entomol. Ital. 22: 38-80.

• Emery, C. 1892a. Sopra alcune formiche raccolte dall'Ingegnere L. Bricchetti Robecchi nel paese dei Somali. [part]. Ann. Mus. Civ. Stor. Nat. 32[=(2(12): 110-112.

• Emery, C. 1893a [1892]. [Untitled. Introduced by: "M. C. Emery, de Bologne, envoie les diagnoses de cinq nouveaux genres de Formicides".]. Bull. Bimens. Soc. Entomol. Fr. 1892:cclxxv-cclxxvii.

• Emery, C. 1893g. Formicides de l'Archipel Malais. Rev. Suisse Zool. 1: 187-229.

• Emery, C. 1893i. Voyage de M. E. Simon aux îles Philippines (mars et avril 1890). Formicides. Ann. Soc. Entomol. Fr. 62: 259-270 PDF'

• Emery, C. 1895n. Sopra alcune formiche della fauna mediterranea. Mem. R. Accad. Sci. Ist. Bologna (5)5:59-75 [pagination of s separate: 59-75.

• Emery, C. 1895m. Viaggio di Leonardo Fea in Birmania e regioni vicine. LXIII. Formiche di Birmania del Tenasserim e dei Monti Carin raccolte da L. Fea. Parte II. Ann. Mus. Civ. Stor. Nat. 34[=(2(14): 450-483.

• Emery, C. 1897c. Formicidarum species novae vel minus cognitae in collectione Musaei Nationalis Hungarici quas in Nova-Guinea, colonia germanica, collegit L. Biró. Természetr. Füz. 20: 571-599.

• Emery, C. 1897e. Viaggio di Lamberto Loria nella Papuasia orientale. XVIII. Formiche raccolte nella Nuova Guinea dal Dott. Lamberto Loria. [concl.]. Ann. Mus. Civ. Stor. Nat. 38[=(2(18): 577-594.

• Emery C (1898) Descrizioni di formiche nuove malesi e australiane. Note sinonimiche. Rendiconti delle Sessioni della Reale Accademia delle Scienze dell’Instituto di Bologna (ns) 2: 231–245.

• Emery C (1899) Formiche di Madagascar raccolte dal Sig. A. Mocquerys nei pressi della Baia di Antongil (1897–1898). Bullettino della Società Entomologica Italiana 31: 263–290.

• Emery C (1900a) Formicidarum species novae vel minus cognitae in collectione Musaei Nationalis Hungarici quas in Nova–Guinea, colonia germanica, collegit L. Biró. Publicatio secunda. Természetajzi Füzetek 23: 310–338.

• Emery C (1900b) Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei. Annali del Museo Civico di Storia Naturale 40[=(2)20]: 661–722.

• Emery C (1901) Ameisen gesammelt in Ceylon von Dr. W. Horn 1899. Deutsche Entomologische Zeitschrift 1901: 113–122.

• Emery C (1906) Note sur Prenolepis vividula Nyl. et sur la classification des espèces du genre Prenolepis. Annales de la Société Entomologique de Belgique 50: 130–134.

• Emery C (1913a) Hymenoptera. Fam. Formicidae. Subfam. Dolichoderinae. Genera Insectorum 137, 50 pp.

• Emery C (1913b) Études sur les Myrmicinae. [V-VII]. Annales de la Société Entomologique de Belgique 57: 250–262.

• Emery C (1923) Einige exotische Ameisen des Deutschen Entomologischen Institutes. Entomologische Mitteilungen 12: 60–62.

• Emery C (1925b) Les espèces européennes et orientales du genre Bothriomyrmex. Bulletin de la Societe Vaudoise des Sciences Naturelles 56: 5–22.

• Erichson WF (1842) Beitrag zur Insecten–Fauna von Vandiemensland, mit besonderer Berücksichtigung der geographischen Verbreitung der Insecten. Archiv für Naturgeschichte 8(1): 83–287.

• Ettershank G (1966) A generic revision of the world Myrmicinae related to Solenopsis and Pheidologeton (Hymenoptera: Formicidae). Australian Journal of Zoology 14: 73–171.

• Evans HE (1985) The pleasures of entomology: portraits of insects and the people who study them. Smithsonian Institution Press, Washington, D.C. 238 pp.

• Fabricius JC (1775) Systema entomologiae, sistens insectorum classes, ordines, genera, species adiectis synonymis, locis, descriptionibus, observationibus. Korte, Flensburgi et Lipsiae [= Flensburg and Leipzig], 832 pp.

• Fabricius JC (1787) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus, genericis, differentiis, specificis, emendationibus, observationibus. Tome 1. C. G. Proft, Hafniae [= Copenhagen], 348 pp.

• Fabricius JC (1793) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tome 2. C. G. Proft, Hafniae [= Copenhagen], 519 pp.

• Fabricius JC (1804) Systema Piezatorum secundum ordines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus. C. Reichard, Brunswick, xiv + 15–439 + 30 pp.

• Fernandez FC (2003) A new myrmicine ant genus from Malaysia with uncertain affinities (Hymenoptera: Formicidae). Zootaxa 341: 1–6.

• Fernandez FC (2004) The American species of the myrmicine ant genus Carebara Westwood (Hymenoptera: Formicidae). Caldasia 26: 191–238.

• Fernandez FC (2010) A new species of Carebara from the Philippines with notes and comments on the systematics of the Carebara genus group (Hymenoptera: Formicidae: Myrmicinae). Caldasia 32(1): 191–203.

• Fisher BL, Bolton B (2007) The ant genus Pseudaphomomyrmex Wheeler, 1920 a junior synonym of Tapinoma Foerster, 1850. Zootaxa 1427: 65–68.

• Foerster A (1850) Hymenopterologische Studien. 1. Formicariae. Ernst Ter Meer, Aachen, 74 pp.

• Forel A (1886a) Indian ants of the Indian Museum, Calcutta, No. 2. Journal of the Asiatic Society of Bengal. Part II. Natural Science 55: 239–249.

• Forel A (1886b) Études myrmécologiques en 1886. Annales de la Société Entomologique de Belgique 30: 131–215.

• Forel A (1890) Fourmis de Tunisie et de l'Algérie orientale. Annales de la Société Entomologique de Belgique 34: lxi–lxxvi.

• Forel A (1891) Les Formicides. [part]. In: Grandidier, A. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (28e fascicule). Hachette et Cie, Paris, v + 237 pp.

• Forel A (1892a) Les Formicides de l'Empire des Indes et de Ceylan. Part I. Journal of the Bombay Natural History Society 7 : 219–245.

• Forel A (1892b) Les Formicides. [concl]. In: Grandidier, A. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (Supplèment au 28e fascicule). Hachette et Cie, Paris, 229–280.

• Forel A (1893a) Les Formicides de l'Empire des Indes et de Ceylan. Part III. Journal of the Bombay Natural History Society 8: 17–36.

• Forel A (1893b) Note préventive sur un nouveau genre et une nouvelle espèce de Formicide (Camponotide). Annales de la Société Entomologique de Belgique 37: 607–608.

• Forel A (1895) [Untitled. Ponera gleadowi n.sp.]. Pp. 60-61 In: Emery, C. Sopre alcune formiche della fauna mediterranea. Memorie della Reale Accademia delle Scienze dell’Istituto di Bologna (5)5: 59–75 [pagination of separate: 292-293].

• Forel A (1900a) Un nouveau genre et une nouvelle espèce de Myrmicide. Annales de la Société Entomologique de Belgique 44: 24–26.

• Forel A (1900b) Ponerinae et Dorylinae d'Australie récoltés par MM. Turner, Froggatt, Nugent, Chase, Rothney, J.–J. Walker, etc. Annales de la Société Entomologique de Belgique 44: 54–77.

• Forel A (1900c) Les Formicides de l'Empire des Indes et de Ceylan. Part VI. Journal of the Bombay Natural History Society 13: 52–65.

• Forel A (1900d) Les Formicides de l'Empire des Indes et de Ceylan. Part VII. Journal of the Bombay Natural History Society 13: 303–332.

• Forel A (1901) Formiciden aus dem Bismarck–Archipel, auf Grundlage des von Prof. Dr. F. Dahl gesammelten Materials bearbeitet. Mitteilungen aus dem Zoologischen Museum in Berlin 2: 1–38.

• Forel A (1902a) Myrmicinae nouveaux de l'Inde et de Ceylan. Revue Suisse de Zoologie 10: 165–249.

• Forel A (1902b) Variétés myrmécologiques. Annales de la Société Entomologique de Belgique 46: 284–296.

• Forel A (1902c) Fourmis nouvelles d'Australie. Revue Suisse de Zoologie 10: 405–548.

• Forel A (1905) Ameisen aus Java. Gesammelt von Prof. Karl Kraepelin 1904. Mitteilungen aus dem Naturalhistorischen Museum in Hamburg 22: 1–26.

• Forel A (1908) Ameisen aus Sao Paulo (Brasilien), Paraguay, etc. Gesammelt von Prof. Herm. v. Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der k.k. Zoologisch-Botanischen Gesellschaft in Wien 58: 340–418.

• Forel A (1909) Ameisen aus Java und Krakatau beobachtet und gesammelt von Herrn Edward Jacobson. Notes from the Leyden Museum 31: 221–232.

• Forel A (1910) Fourmis des Philippines. Philippine Journal of Science Section D. General Biology, Ethnology, and Anthropology 5: 121–130.

• Forel A (1911a) Fourmis de Bornéo, Singapore, Ceylan, etc. récoltées par MM. Haviland, Green, Winkler, Will, Hose, Roepke et Waldo. Revue Suisse de Zoologie 19: 23–62.

• Forel A (1911b) Ameisen aus Java beobachtet und gesammelt von Herrn Edward Jacobson. II. Theil. Notes from the Leyden Museum 33: 193–218.

• Forel A (1911c) Sur le genre Metapone n. g. nouveau groupe des Formicides et sur quelques autres formes nouvelles. Revue Suisse de Zoologie 19: 445–459.

• Forel A (1912a) Ameisen aus Java beobachtet und gesammelt von Edward Jacobson. III. Theil. Notes from the Leyden Museum 34: 97–112.

• Forel A (1912b) Einige neue und interessante Ameisenformen aus Sumatra etc. Zoologische Jahrbücher. Supplement 15 ("Erster Band”): 51–78.

• Forel A (1913) Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien ausgeführt im Auftrage der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel–Reepen. II. Ameisen aus Sumatra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr. v. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 36: 1–148.

• Forel A (1915) Fauna Simalurensis. Hymenoptera Aculeata, Fam. Formicidae. Tijdschrift voor Entomologie 58: 22–43.

• General, D.E.M. 2017. First Philippine record of the Neotropical ant, Strumigenys eggersi Emery (Hymenoptera:Formicidae). Philipp Ent 31(1):73-78.

• General, D.M. & Alpert, G.D. 2012. A synoptic review of the ant genera of the Philippines. ZooKeys 200: 1-111.

• Gobin B, Peeters C, Billen J (1998) Colony reproduction and arboreal life in the ponerine ant Gnamptogenys menadensis (Hymenoptera, Formicidae). Netherlands Journal of Zoology 48(1): 53–63.

• Gobin B, Billen J, Peeters C (1999) Policing behaviour towards virgin egg layers in a polygynous ponerine ant. Animal Behaviour 58: 1117–1122.

• Gregg RE (1951) Two new species of exotic ants. Psyche (Cambridge) 58: 77–84.

• Heaney LR, Balete DS, Ingle N, Lepiten M, Oliver W, Rickart EA, Utzurrum RCB, Gonzales PC (1999) Mammalian diversity on Mount Isarog, a threatened ceneter of endemism on southern Luzon Island, Philippines. Fieldiana Zoology new series 95: 1–62.

• Heinze J, Kühnholz S, Schilder K, Hölldobler B (1993) Behavior of ergatoid males in the ant, Cardiocondyla nuda. Insectes Sociaux 40: 273–282.

• Heterick BE (2001) Revision of the Australian ants of the genus Monomorium (Hymenoptera, Formicidae). Invertebrate Taxonomy 15: 353–459.

• Hosoishi S, Ogata K (2009a) A taxonomic revision of the Asian endemic subgenus Physocrema of the genus Crematogaster (Hymenoptera: Formicidae). Zootaxa 2062: 15–36.

• Hosoishi S, Ogata K (2009b) A checklist of the ant genus Crematogaster in Asia (Hymenoptera: Formicidae). Bulletin of the Institute of Tropical Agriculture, Kyushu University 32: 43–83.

• Hung ACF (1967) A revision of the ant genus Polyrhachis at the subgeneric level. Transactions of the American Entomological Society 93: 395–422.

• International Organization for Standardization (2010) English country names and code elements.. Available at: http://www.iso.org/iso/country_codes/iso_3166_code_lists/english_country_names_and_code_elements.htm [accessed 28.X.2010].

• Jerdon TC (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science 17: 103–127.

• Karavaiev V (1926) Über den Nestbau von Polyrhachis (subg. Myrmhopla) tubifex sp. n. (Fam. Formicidae). Biologisches Zentralblatt 46: 143–145.

• Karavaiev V (1935) Neue ameisen aus dem Indo–Australischen Gebiet, nebst Revision einiger Formen. Treubia 15: 57–118.

• Kohout RJ (1987) Three new Polyrhachis sexspinosa–group species from the Philippines. Memoirs of the Queensland Museum 25: 169–176.

• Kohout RJ (1994) Polyrhachis lama, a new ant from the Tibetan plateau (Formicidae, Formicinae). Memoirs of the Queensland Museum 35: 137–138.

• Kohout RJ (1998) New synonyms and nomenclatural changes in the ant genus Polyrhachis Fr. Smith (Hymenoptera, Formicidae, Formicinae) Memoirs of the Queensland Museum 42(2): 505–531.

• Kohout RJ (2006a) A review of the Polyrhachis cryptoceroides species-group with description of a new species from Thailand (Hymenoptera: Formicidae) Myrmecologische Nachrichten 8: 145–150.

• Kohout RJ (2006b) New species of the Polyrhachis (Myrma) parabiotica species group from the Philippines. Australian Entomologist 33: 155–163.

• Kutter H (1931) Forelophilus, eine neue Ameisengattung. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 15: 193–194.

• LaPolla JS (2004) Acropyga (Hymenoptera, Formicidae) of the World. Contributions of the American Entomological Institute 33(3): 1–130.

• LaPolla JS (2009) Taxonomic revision of the Southeast Asian ant genus Euprenolepis. Zootaxa 2046: 1–25.

• LaPolla JS, Brady SG, Shattuck SO (2010) Phylogeny and taxonomy of the Prenolepis genus-group of ants (Hymenoptera: Formicidae). Systematic Entomology 35: 118–131.

• LaPolla JS, Longino JT (2006) An unusual new Brachymyrmex Mayr (Hymenoptera: Formicidae) from Costa Rica, with implications for the phylogeny of the Lasiine tribe group. Proceedings of the Entomological Society of Washington 108(2): 297–305.

• Latreille PA (1802) Histoire naturelle générale et particulière des Crustacés et des insectes. Tome 3. Familles naturelles des genres. F. Dufart, Paris, xii + 467 pp.

• Latreille PA (1804) Tableau méthodique des insectes. pp. 129–200 In: Société de Naturalistes et d'Agriculteurs. Nouveau dictionnaire d'histoire naturelle. Tome 24. Déterville, Paris, 84 + 85 + 238 + 18 + 34 pp.

• Lattke JE (2004) A taxonomic revision and phylogenetic analysis of the ant genus Gnamptogenys Roger in Southeast Asia and Australasia (Hymenoptera, Formicidae, Ponerinae). University of California Publications in Entomology 122: 1–266.

• Le Guillou EJF (1842) Catalogue raisonné des insectes hyménoptères recueillis dans le voyage de circumnavigation des corvettes l'Astrolabe et la Zélée. Annales de la Société Entomologique de France 10: 311–324.

• Linnaeus C (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. L. Salvii, Holmiae [= Stockholm], 824 pp.

• Lucky A, Ward PS (2010) Taxonomic revision of the ant genus Leptomyrmex Mayr (Formicidae: Dolichoderinae). Zootaxa 2688: 1–67.

• Lund AW (1831) Lettre sur les habitudes de quelques fourmis du Brésil, adressée à M. Audouin. Annales des Sciences Naturelles 23: 113–138.

• Mackay WP, Mackay EE (2010) The systematics and biology of the New World ants of the genus Pachycondyla. Edwin Mellen Press, 642 p.

• Mann WM (1919) The ants of the British Solomon Islands. Bulletin of the Museum of Comparative Zoology 63: 273–391.

• Mann WM (1921) The ants of the Fiji Islands. Bulletin of the Museum of Comparative Zoology 64: 401–499.

• Mayr G (1853) Beiträge zur Kenntniss der Ameisen. Verhandlungen der Zoologisch–Botanischen Vereins in Wien 3: 101–114.

• Mayr G (1855) Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate aufgefundenen Ameisen, nebst Hinzufügung jener in Deutschland, in der Schweiz und in Italien vorkommenden Arten. Verhandlungen der Zoologisch–Botanischen Vereins in Wien 5: 273–478.

• Mayr G (1861) Die Europäischen Formiciden. Nach der analytischen Methode bearbeitet. C. Gerolds Sohn, Wien, 80 pp.

• Mayr G (1862) Myrmecologische Studien. Verhandlungen der Kaiserlich–Königlichen Zoologisch–Botanischen Geselschaft in Wien 12: 649–776.

• Mayr G (1865) Formicidae. In: Reise der Österreichischen Fregatte "Novara" um die Erde in den Jahren 1857, 1858, 1859, unter den befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Bd. II. Abt. 1. Wien, K. Gerold's Sohn, 119 pp.

• Mayr G (1866a) Myrmecologische Beiträge. Sitzungberichte der Kaiserlichen Akademie der Wissenschaften in Wien. Mathematisch–Naturwissenschaftliche Classe. Abteilung I. 53: 484–517.

• Mayr G (1866b) Diagnosen neuer und wenig gekannter Formiciden. Verhandlungen der Kaiserlich–Königlichen Zoologisch–Botanischen Geselschaft in Wien 16: 885–908.

• Mayr G (1867) Adnotationes in monographiam formicidarum Indo–Neerlandicarum. Tijdschrift voor Entomologie 10: 33–117.

• Mayr G (1868) Die Ameisen des baltischen Bernsteins. Beiträge zur Naturkunde Preussens 1: 1–102.

• Mayr G (1870) Neue Formiciden. Verhandlungen der Kaiserlich–Königlichen Zoologisch–Botanischen Geselschaft in Wien 20: 939–996.

• Mayr G (1872) Formicidae Borneenses collectae a J. Doria et O. Beccari in territorio Sarawak annis 1865–1867 Annali del Museo Civico di Storia Naturale 2: 133–155.

• Mayr G (1879) Beiträge zur Ameisen–Fauna Asiens. Verhandlungen der Kaiserlich–Königlichen Zoologisch–Botanischen Geselschaft in Wien 28: 645–686.

• Mayr G (1887) Südamerikanische Formiciden. Verhandlungen der Kaiserlich–Königlichen Zoologisch–Botanischen Geselschaft in Wien 37: 511–632.

• Mayr G (1901) Südafrikanische Formiciden, gesammelt von Dr. Hans Brauns. Annalen des Kaiserlich–Königlichen Naturhistorischen Museums in Wien 16: 1–30.

• Menozzi C (1922) Miscellanea mirmecologica. [concl] Annali del Museo Civico di Storia Naturale "Giacomo Doria" 49[=(3)(9)]: 353–358.

• Menozzi C (1925) Nouvelles fourmis des Philippines. Philippine Journal of Science 28 : 439–451.

• Menozzi C (1926) Nuove formiche delle isole Filippine e di Singapore. Atti della Società dei Naturalisti e Matematici di Modena 56[=(6)4]: 92–103.

• Menozzi C (1932) Una nuova specie de Polyrhachis (Hym.–Formicidae) e descrizione di tre nidi di formiche appartenenti a questo genere. Wiener Entomologische Zeitung 49: 303–308.

• Menozzi C (1933) Description préliminaire d'une espèce nouvelle de fourmi constituant un genre nouveau. Natuurhistorisch Maandblad 22: 146–147.

• Menozzi C (1939) Qualche nuova formica di Sumatra. Tijdschrift voor Entomologie 82: 175–181.

• Mercier J-L, Lenior J-C, Eberhardt A, Frohschammer S, Williams C, Heinze J (2007) Hammering, mauling, and kissing: stereotyped courtship behavior in Cardiocondyla ants. Insectes Sociaux 54: 403–411.

• Moffett MW (1985) Revision of the genus Myrmoteras. Bulletin of the Museum of Comparative Zoology 151: 1–53.

• Moffett MW (1986) Revision of the myrmicine genus Acanthomyrmex. Bulletin of the Museum of Comparative Zoology 151: 55–89.

• Motschoulsky V de (1863) Essai d'un catalogue des insectes de l'île Ceylan (suite). Bulletin de la Société Impériale des Naturalistes de Moscou 36(3): 1–153.

• Naskrecki, P. (2004) The taxonomic impediment, a colonial legacy. Wings (Fall 2004) 27(2): 20–24.

• Nylander W (1846) Additamentum adnotationum in monographiam formicarum borealium Europae. Acta Societatis Scientiarum Fennicae 2: 1041–1062.

• Ogata K, Terayama M, Masuko K (1995) The ant genus Leptanilla, discovery of the worker–associated male of L. japonica, and a description of a new species from Taiwan. Systematic Entomology 20: 27–34.

• Philippine Atmospheric, Geophysical and Astronomical Services Administration (2010). PAGASA. Available at: http://www.pagasa.dost.gov.ph/ [accessed 27.IV.2010].

• Peeters C, Higashi S (1989) Reproductive dominance controlled by mutilation in the queenless ant Diacamma australe. Naturwissenschaften 76: 177–180.

• Petersen B (1968) Some novelties in presumed males of Leptanillinae (Hymenoptera, Formicidae). Entomologiske Meddelelser 36: 577–598.

• Rigato F (1994) Revision of the myrmicine ant genus Lophomyrmex, with a review of its taxonomic position (Hymenoptera: Formicidae). Systematic Entomology 19: 47–60.

• Rigato F, Bolton B (2001) The ant genus Liomyrmex, a review. Bollettino della Societa Entomologica Italiana 133: 247–256.

• Roger J (1859) Beiträge zur Kenntniss der Ameisenfauna der Mittelmeerländer. I. Berliner Entomologische Zeitschrift 3: 225–259.

• Roger J (1860) Die Ponera–artigen Ameisen. Berliner Entomologische Zeitschrift 4: 278–312.

• Roger J (1861) Die Ponera–artigen Ameisen (Schluss). Berliner Entomologische Zeitschrift 5: 1–54.

• Roger J (1862) Einige neue exotische Ameisen–Gattungen und Arten. Berliner Entomologische Zeitschrift 6: 233–254.

• Roger J (1863) Die neu aufgeführten Gattungen und Arten meines Formiciden–Verzeichnisses nebst Ergänzung einiger früher gegebenen Beschreibungen. Berliner Entomologische Zeitschrift 7: 131–214.

• Samson SA, Rickart EA, Gonzales PC (1997) Ant diversity and abundance along an elevational gradient in the Philippines. Biotropica 29: 349–363.

• Santschi F (1910) Deux nouvelles fourmis du Tonkin. Naturaliste 32: 283–284.

• Santschi F (1914) Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale, 1911–1912. Résultats scientifiques. Insectes Hyménoptères. II. Formicidae. Libr. A. Schulz, Paris, 41–148.

• Santschi F (1925) Revision des Myrmicaria d’Afrique. Annales de la Société Entomologique de Belgique 64 (1924): 133–176.

• Santschi F (1926) Trois notes myrmécologiques. Annales de la Société Entomologique de France 95: 13–28.

• Santschi F (1928) Nouvelles fourmis de Chine et du Turkestan Russe. Bulletin et Annales de la Société Entomologique de Belgique 68: 31–46.

• Santschi F (1938) Notes sur quelques Ponera Latr. Bulletin de la Société Entomologique de France 43: 78–80.

• Saunders WW (1842) Descriptions of two hymenopterous insects from northern India. Transactions of the Entomological Society of London 3: 57–58.

• Schneirla TC, Reyes AY (1966) Raiding and related behavior in two surface–adapted species of the Old World Doryline ant, Aenictus. Animal Behavior 14: 132–148.

• Schlick-Steiner BC, Steiner FM, Zettel H (2006) Tetramorium pacificum Mayr, 1870, T. scabrum Mayr, 1879 sp. rev., T. manobo (Calilung, 2000) (Hymenoptera: Formicidae) – three good species. Myrmecologische Nachrichten 8: 181-191.

• Schödl S (1998) Taxonomic revision of Oriental Meranoplus F. Smith, 1853 (Insecta, Hymenoptera, Formicidae, Myrmicinae). Annalen des Naturhistorischen Museums in Wien 100: 361–394.

• Seifert B (2003) The ant genus Cardiocondyla (Insecta: Hymenoptera: Formicidae) - a taxonomic revision of the C. elegans, C. bulgarica, C. batesii, C. nuda, C. shuckardi, C. stambuloffii, C. wroughtonii, C. emeryi and C. minutior species groups. Annalen des Naturhistorischen Museums in Wien 104(B): 203-338.

• Shattuck SO (1992a) Review of the dolichoderine ant genus Iridomyrmex Mayr with descriptions of three new genera (Hymenoptera, Formicidae). Journal of the Australian Entomological Society 31: 13–18.

• Shattuck SO (1992b) Higher classification of the ant subfamilies Aneuretinae, Dolichoderinae and Formicinae (Hymenoptera, Formicidae). Systematic Entomology 17: 199–206.

• Shattuck SO (1992c) Generic revision of the ant subfamily Dolichoderinae (Hymenoptera, Formicidae). Sociobiology 21: 1–181.

• Shattuck SO (1999) Australian ants, their biology and identification. Monographs on Invertebrate Taxonomy Vol. 3, CSIRO Publishing, Collingwood, Victoria, Australia. xi + 226 pp.

• Shattuck SO (2008a) Revision of the ant genus Prionopelta (Hymenoptera, Formicidae) in the Indo-Pacific region. Zootaxa 1846: 21–34.

• Shattuck SO (2008b) Review of the ant genus Aenictus (Hymenoptera: Formicidae) in Australia with notes on A. ceylonicus (Mayr). Zootaxa 1926: 1–19.

• Shattuck SO (2011) Revision of the ant genus Calyptomyrmex (Hymenoptera: Formicidae) in South-east Asia and Oceania. Zootaxa 2743: 1–26.

• Shattuck SO, Barnett NJ (2007) Revision of the ant genus Mayriella (Hymenoptera, Formicidae). In: Snelling R R, Fisher BL, Ward PS (Eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute 80: 437–458.

• Shattuck SO, Barnett NJ (2010) Ants Down Under, CSIRO, Australia. Available at: http://anic.ento.csiro.au/ants [accessed 4.IV.2010].

• Shuckard WE (1840) Monograph of the Dorylidae, a family of the Hymenoptera Heterogyna. (Continued from p. 201). Annals of Natural History 5: 258–271.

• Smith F (1851) List of the specimens of British animals in the collection of the British Museum. Part VI. Hymenoptera, Aculeata. British Museum, London, 134 pp.

• Smith F (1852) Descriptions of some hymenopterous insects captured in India, with notes on their economy, by Ezra T. Downes, Esq., who presented them to the Honourable the East India Company. Annals and Magazine of Natural History 2(9): 44–50.

• Smith F (1853) Monograph of the genus Cryptocerus, belonging to the group Cryptoceridae – family Myrmicidae – division Hymenoptera Heterogyna. Transactions of the Entomological Society of London 2: 213–228.

• Smith F (1857) Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and at Singapore, by A. R. Wallace. [part]. Journal and Proceedings of the Linnean Society of London. Zoology 2: 42–88.

• Smith F (1858) Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. British Museum, London, 216 pp.

• Smith F (1859) Catalogue of hymenopterous insects collected by Mr. A. R. Wallace at the islands of Aru and Key. [part]. Journal and Proceedings of the Linnean Society of London. Zoology 3: 132–158.

• Smith F (1860a) Descriptions of new species of hymenopterous insects collected by Mr. A. R. Wallace at Celebes. Journal of the Proceedings of the Linnean Society of London. Zoology 5(17b)(supplement to vol. 4) 4: 57–93.

• Smith F (1860b) Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the islands of Bachian, Kaisaa, Amboyna, Gilolo, and at Dory in New Guinea. Journal of the Proceedings of the Linnean Society of London. Zoology 5(17b) (supplement to vol. 4) 4: 93–143.

• Smith F (1860c) Descriptions of new genera and species of exotic Hymenoptera. Journal of Entomology 1: 65–84.

• Smith F (1861) Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the islands of Ceram, Celebes, Ternate, and Gilolo. [part]. Journal of the Proceedings of the Linnean Society of London. Zoology 6: 36–48.

• Smith F (1865) Descriptions of new species of hymenopterous insects from the islands of Sumatra, Sula, Gilolo, Salwatty, and New Guinea, collected by Mr. A. R. Wallace. Journal of the Proceedings of the Linnean Society of London. Zoology 8: 61–94.

• Smith F (1876) Descriptions of three new species of Hymenoptera (Formicidae) from New Zealand. Transactions of the Entomological Society of London 1876: 489–492.

• Smith F (1877) Descriptions of new species of the genera Pseudomyrma and Tetraponera, belonging to the family Myrmicidae. Transactions of the Entomological Society of London 1877: 57–72.

• Sorger DM, Zettel H (2009) Polyrhachis (Myrma) cyaniventris F. Smith, 1858 (Hymenoptera: Formicidae) and a related new ant species from the Philippines. Zootaxa 2174: 27–37.

• Sorger DM, Zettel H (2010) Taxonomic additions to the Polyrhachis (Myrma) cyaniventris species group (Insecta: Hymenoptera: Formicidae). Annalen des Naturhistorischen Museums in Wien, B 111: 31–36.

• Sorger DM, Zettel H (2011) On the ants (Hymenoptera: Formicidae) of the Philippine Islands: V. The genus Odontomachus Latreille, 1804. Myrmecological News 14: 141–163.

• Stitz H (1925) Ameisen von den Philippinen, den malayischen und ozeanischen Inseln. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1923: 110–136.

• Stitz H (1938) Neue Ameisen aus dem indo–malayischen Gebiet. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1938: 99–122.

• Taylor RW (1965) A monographic revision of the rare tropicopolitan ant genus Probolomyrmex Mayr. Transactions of the Royal Entomological Society of London 117: 345–365.

• Taylor RW (1966) Notes on the Indo–Australian ants of genus Simopone Forel (Hymenoptera–Formicidae). Psyche (Cambridge) 72: 287–290.

• Taylor RW (1967) A monographic revision of the ant genus Ponera Latreille. Pacific Insects Monograph 13: 1–112.

• Taylor RW (1968) Notes on the Indo–Australian basicerotine ants. Australian Journal of Zoology 16: 333–348.

• Taylor RW (1979) Melanesian ants of the genus Amblyopone. Australian Journal of Zoology 26 (1978): 823–839.

• Taylor RW (1980) Australian and Melanesian ants of the genus Eurhopalothrix Brown and Kempf – notes and new species. Australian Journal of Zoology 27(1979): 773–788.

• Taylor RW (1985) The ants of the Papuasian genus Dacetinops (Hymenoptera, Formicidae, Myrmicinae). In: Ball GE (Ed.) Taxonomy, phylogeny and zoogeography of beetles and ants, a volume dedicated to the memory of Philip Jackson Darlington, Jr., 1904–1983. Series Entomologica (Dordrecht) 33: 41-67.

• Taylor RW (1990) New Asian ants of the tribe Basicerotini, with an on–line computer interactive key to the twenty–six known Indo–Australian species (Hymenoptera, Formicidae, Myrmicinae). Invertebrate Taxonomy 4: 397–425.

• Taylor RW (1991a) Notes on the ant genera Romblonella and Willowsiella, with comments on their affinities, and the first description of Australian species. Psyche (Cambridge) 97: 281–296.

• Terayama M (1985) Two new species of the genus Acropyga (Hymenoptera: Formicidae) from Taiwan and Japan. Kontyû 53: 284-289.

• Terayama M (1999) Taxonomic studies of the Japanese Formicidae. Part 6. Genus Cardiocondyla Emery. Memoirs of the Myrmecological Society of Japan 1: 99–107.

• Terayama M, Ito F, Gobin B (1998) Three new species of the genus Acanthomyrmex Emery (Hymenoptera, Formicidae) from Indonesia, with notes on the reproductive caste and colony composition. Entomological Science 1(2): 257–264.

• Terayama M, Yamane S (1989) The army ant genus Aenictus from Sumatra, with descriptions of three new species. Japanese Journal of Entomology 57: 597–603.

• Trager JC (1991) A revision of the fire ants, Solenopsis geminata group. Journal of the New York Entomological Society 99: 141–198.

• Van Steenis-Kruseman, M.J.(2006). Cyclopedia of Malesian collectors. Available at: http://www.nationaalherbarium.nl/fmcollectors/Home.htm [12.II.2011].

• Viehmeyer H (1913) Ameisen aus dem Kopal von Celebes. Stettiner Entomologische Zeitung 74: 141–155.

• Viehmeyer H (1916a) Ameisen von Singapore. Beobachtet und gesammelt von H. Overbeck. Archiv für Naturgeschichte(A) 81(8): 108–168.

• Viehmeyer H (1916b) Ameisen von den Philippinen und anderer Herkunft (Hym.). Entomologische Mitteilungen 5: 283–291.

• Viehmeyer H (1922) Neue Ameisen. Archiv für Naturgeschichte (A) 88(7): 203–220.

• Walker F (1859) Characters of some apparently undescribed Ceylon insects. [part]. Annals and Magazine of Natural History 3(4): 370–376.

• Wang M (2003) A monographic revision of the ant genus Pristomyrmex (Hymenoptera, Formicidae). Bulletin of the Museum of Comparative Zoology 157(6): 383–542.

• Ward PS (2001) Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera, Formicidae) in the Oriental and Australian regions. Invertebrate Taxonomy 15: 589–665.

• Ward PS, Bolton B, Shattuck SO, Brown WL Jr (1996) A bibliography of ant systematics. University of California Publications in Entomology 116: 1–417.

• Westwood JO (1839) An introduction to the modern classification of insects; founded on the natural habits and corresponding organisation of the different families. Volume 2. Part XI. Longman, Orme, Brown, Green and Longmans, London, pp. 193–224.

• Westwood JO (1840) Observations on the genus Typhlopone, with descriptions of several exotic species of ants. Annals and Magazine of Natural History 6: 81–89.

• Wheeler WM (1909) Ants of Formosa and the Philippines. Bulletin of the American Museum of Natural History 26: 333–345.

• Wheeler WM (1916) Four new and interesting ants from the mountains of Borneo and Luzon. Proceedings of the New England Zoological Club 6: 9–18.

• Wheeler WM (1917) A new Malayan ant of the genus Prodiscothyrea. Psyche (Cambridge) 24: 29–30.

• Wheeler WM (1919a) The ants of Borneo. Bulletin of the Museum of Comparative Zoology 63: 43–147.

• Wheeler WM (1919b) The ant genus Lordomyrma. Psyche (Cambridge) 26: 97–106.

• Wheeler WM (1919c) The ants of the genus Metapone Forel. Annals of the Entomological Society of America 12: 173–191.

• Wheeler WM (1924a) Ants of Krakatau and other islands in the Sunda Strait. Treubia 5: 239–258.

• Wheeler WM (1924b) On the ant genus Chrysapace Crawley. Psyche (Cambridge) 31: 224–225

• Wheeler WM (1927) The physiognomy of insects. Quarterly Review of Biology 2: 1–36.

• Wheeler WM (1928) A new species of Probolomyrmex from Java. Psyche (Cambridge) 35: 7–9.

• Wheeler WM (1929a) The identity of the ant genera Gesomyrmex Mayr and Dimorphomyrmex Ernest André. Psyche (Cambridge) 36: 1–12.

• Wheeler WM (1929b) Note on Gesomyrmex. Psyche (Cambridge) 36: 91–92.

• Wheeler WM (1929c) The ant genus Rhopalomastix. Psyche (Cambridge) 36: 95–101.

• Wheeler WM (1929d) Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Bollettino del Laboratorio di Zoologia generale e agrarian del R. Instituto Superiore agrario di Portici 24: 27–64.

• Wheeler WM (1930a) A second note on Gesomyrmex. Psyche (Cambridge) 37: 35–40.

• Wheeler WM (1930b) Two new genera of ants from Australia and the Philippines. Psyche (Cambridge) 37: 41–47.

• Wheeler WM (1930c) Philippine ants of the genus Aenictus with descriptions of the females of two species. Journal of the New York Entomological Society 38: 193–212.

• Wheeler WM (1933) Three obscure genera of ponerine ants. American Museum Novitiates 672: 1–23.

• Wheeler WM (1934) A second revision of the ants of the genus Leptomyrmex Mayr. Bulletin of the Museum of Comparative Zoology 77: 69–118.

• Wheeler WM (1935a) Two new genera of myrmicine ants from Papua and the Philippines. Proceedings of the New England Zoological Club 15: 1–9.

• Wheeler WM (1935b) New ants from the Philippines. Psyche (Cambridge) 42: 38–52.

• Wheeler WM, Chapman JW (1925) The ants of the Philippine Islands. Part I, Dorylinae and Ponerinae. Philippine Journal of Science 28: 47–73.

• Willey RB, Brown WL Jr (1983) New species of the ant genus Myopias. Psyche (Cambridge) 90: 249–285.

• Wilson EO (1958) Studies on the ant fauna of Melanesia. 1. The tribe Leptogenyini. 2. The tribes Amblyoponini and Platythyreini. Bulletin of the Museum of Comparative Zoology 118: 101–153.

• Wilson EO (1964) The true army ants of the Indo–Australian area. Pacific Insects 6: 427–483.

• WitteV, Maschwitz U (2008) Mushroom harvesting ants in the tropical rain forest. Naturwissenschaften 95(11): 1049–54.

• Yamane S (2009) Odontoponera denticulata (F. Smith) (Formicidae: Ponerinae), a distinct species inhabiting disturbed areas. Ari, Journal of the Myrmecological Society of Japan 32: 1–8.

• Yamauchi K, Asano Y, Lautenschläger B, Trindl A, Heinze J (2005) A new type of male dimorphism with ergatoid and short-winged males in Cardiocondyla cf. kagutsuchi. Insectes Sociaux 52: 274–281.

• Yanoviak SP, Fisher BL, Alonso A (2008) Directed aerial descent behavior in African canopy ants (Hymenoptera: Formicidae). Journal of Insect Behavior 21: 164–171.

• Zettel, H. 2006. On the ants of the Philippine Islands: I. The genus Pristomyrmex Mayr, 1866. Myrmecologische Nachrichten 8:59-68.

• Zettel H (2007) A new species of Pristomyrmex Mayr 1866 (Hymenoptera: Formicidae) from Cebu, the Philippines. Linzer Biologische Beiträge 39(2): 1251–1255.

• Zettel H (2008) On the ants of the Philippine Islands: III. The genus Recurvidris Bolton. Linzer Biologische Beiträge 40(1): 891–895.

• Zettel H and Sorger, D.M. (2010a) On the ants (Hymenoptera: Formicidae) of the Philippine Islands: IV. The genus Vombisidris Bolton 1991. Entomologica Austriaca 17: 37–44.

• Zettel H, Sorger DM (2010b) Three new species of the army ant genus Aenictus Shuckard, 1840 (Hymenoptera: Formicidae: Aenictinae) from Borneo and the Philippines. Zeitschrift Arbeitsgemeinschaft Österreichischer Entomologen 62: 115–125.

• Zettel, H. & Sorger, D.M. 2011. New Myrmoteras ants from the southeastern Philippines. Raffles Bulletin of Zoology 59: 61-67.

• Zettel H, Zimmerman D (2007) On the ants (Hymenoptera: Formicidae) of the Philippine Islands: II. The genus Forelophilus Kutter, 1931. Asian Myrmecology (Taxonomy) 1: 19–30.

Figure legends Figure 1. Full-face and profile images of Philippine ant genera. Acanthomyrmex mindanao Moffett, 1986, major worker (A, B). Acropyga pallida (Donisthorpe, 1938), minor worker (C, D). Aenictus ceylonicus (Mayr, 1866) (E, F). Figure 2. Full-face and profile images of Philippine ant genera. Amblyopone luzonica (Wheeler and Chapman, 1925) (A, B). Anillomyrma decamera (Emery, 1901) (images reproduced with permission of Dr. Katsuyuki Eguchi) (C, D). Anochetus isolatus Mann, 1919 (E, F). Figure 3. Full-face and profile images of Philippine ant genera. Anoplolepis gracilipes (A, B). Aphaenogaster species PH-02 (C, D). Bothriomyrmex species PH-01 (E, F). Figure 4. Full-face and profile images of Philippine ant genera. Calyptomyrmex beccarii Emery, 1887 (A, B). Camponotus albocinctus (Ashmead, 1905), major worker (C, D). Cardiocondyla sima Wheeler, 1935, queen (E, F). Figure 5. Full-face and profile images of Philippine ant genera. Carebara alperti Fernandez, 2010 (A, B). Cataulacus chapmani Bolton, 1974 (C, D). Centromyrmex feae (Emery, 1889) (E, F). Figure 6. Full-face and profile images of Philippine ant genera. Cerapachys rufithorax Wheeler and Chapman, 1925 (A, B). Crematogaster difformis F. Smith, 1857 (C, D). Dacetinops cirrosus (E, F). Figure 7. Full-face and profile images of Philippine ant genera. Diacamma panayense Wheeler and Chapman, 1925 (A, B). Dilobocondyla chapmani Wheeler, 1924 (C, D). Discothyrea clavicornis Emery, 1897 (E, F). Figure 8. Full-face and profile images of Philippine ant genera. Dolichoderus thoracicus (A, B). Echinopla pallipes F. Smith, 1857 (C, D). Euprenolepis species PH-01 (E, F). Figure 9. Full-face and profile images of Philippine ant genera. Eurhopalothrix philippina Brown and Kempf, 1960 (A, B). Forelophilus species PH-01 (C, D). Gauromyrmex acanthina (Karavaiev, 1935) (E, F). Figure 10. Full-face and profile images of Philippine ant genera. Gesomyrmex luzonensis, major worker (A, B). Gnamptogenys chapmani Brown, 1958 (C, D). Harpegnathos venator Donisthorpe, 1937 (E, F). Figure 11. Full-face and profile images of Philippine ant genera. Hypoponera confinis (Roger, 1860) (A, B). Iridomyrmex species PH-01 (C, D). Lepisiota chapmani Wheeler, 1935 (E, F). Figure 12. Full-face and profile images of Philippine ant genera. Leptanilla astylina (line drawing reprinted from Entomologiske Meddelelser, with permission) (A). Leptogenys maxillosa (F. Smith, 1858) (B, C). Leptomyrmex fragilis (D, E). Figure 13. Full-face and profile images of Philippine ant genera. Liomyrmex gestroi (A, B). Lophomyrmex bedoti (C, D). Lordomyrma species PH-01 (E, F). Figure 14. Full-face and profile images of Philippine ant genera. Mayriella transfuga (A, B). Meranoplus species PH-01 (C, D). Metapone gracilis Wheeler, 1935 (E, F). Figure 15. Full-face and profile images of Philippine ant genera. Monomorium species PH-01 (A, B). Myopias lobosa Willey and Brown, 1983 (C, D). Myopopone castanea (F. Smith 1860) (E, F). Figure 16. Full-face and profile images of Philippine ant genera. Myrmecina species PH-01 (A, B). Myrmicaria brunnea Saunders, 1842 (C, D). Myrmoteras wlliamsi Wheeler, 1919 (E, F). Figure 17. Full-face and profile images of Philippine ant genera. Mystrium camillae Emery, 1889 (A, B). Noonilla copiosa Petersen, 1968 (line drawings reprinted from Entomologiske Meddelelser, with permission) (C, D). Nylanderia species PH-01 (E, F). Figure 18. Full-face and profile images of Philippine ant genera. Ochetellus glaber (A, B). Odontomachus simillimus F. Smith, 1858 (C, D). Odontoponera denticulata (E, F). Figure 19. Full-face and profile images of Philippine ant genera. Oecophylla smaragdina (Fabricius, 1775) (A, B). Overbeckia subclavata Viehmeyer, 1916 (C, D). Pachycondyla claudata (Menozzi, 1926) (E, F). Figure 20. Full-face and profile images of Philippine ant genera. Paraparatrechina iridescens (A, B). Paratopula macta Bolton, 1988 (C, D). Paratrechina longicornis (Latreille, 1802) (E, F). Figure 21. Full-face and profile images of Philippine ant genera. Pheidole hortensis Forel, 1913, major worker (A, B). Pheidologeton maccus Wheeler, 1929, major worker (C, D). Philidris species PH-01 (E, F). Figure 22. Full-face and profile images of Philippine ant genera. Plagiolepis species PH-01 (A, B). Platythyrea parallela (F. Smith, 1859) (C, D). Polyrhachis cyaniventris F. Smith, 1858 (E, F). Figure 23. Full-face and profile images of Philippine ant genera. Ponera oreas (Wheeler, 1933) (A, B). Prenolepis species PH-01 (C, D). Prionopelta kraepelini Forel, 1905 (E, F). Figure 24. Full-face and profile images of Philippine ant genera. Pristomyrmex bicolor Emery, 1900 (A, B). Probolomyrmex dammermani (C, D). Proceratium papuanum (E, F). Figure 25. Full-face and profile images of Philippine ant genera. Pseudolasius typhlops Wheeler, 1935, major worker (A, B). Pyramica pedunculata (Brown, 1953) (C, D). Recurvidris species PH-01 (E, F). Figure 26. Full-face and profile images of Philippine ant genera. Rhopalomastix species PH-01 queen (A, B). Rhoptromyrmex wroughtonii (C, D). Rhytidoponera species PH-01 (E, F). Figure 27. Full-face and profile images of Philippine ant genera. Romblonella opaca (A, B). Simopone conradti Emery, 1899 (N.B. Not from the Philippines, the unique holotype of S. chapmani is badly broken and cannot help the student visualize this rare genus) (C, D). Solenopsis geminata minor worker (E, F). Figure 28. Full-face and profile images of Philippine ant genera. Strumigenys chapmani Brown, 1954 (A, B). Tapinoma williamsi (Wheeler, 1935b) (C, D). Technomyrmex albipes (F. Smith, 1861) (E, F). Figure 29. Full-face and profile images of Philippine ant genera. Tetramorium khnum Bolton, 1977 (A, B). Tetraponera allaborans (Walker, 1859) (C, D). Tyrannomyrmex species PH-01 (E, F). Figure 30. Full-face and profile images of Philippine ant genera. Vollenhovia ambitiosa Menozzi, 1925 (A, B). Vombisidris philippina Zettel and Sorger, 2010 (image reproduced with permission of NHMW and antbase.net) (C, D). Unnamed genus PH-01 (E, F). Figure 31. Full-face and profile images of Philippine ant genera. Unnamed genus PH-02 (A, B). Unnamed genus PH-03 (C, D).

Appendix

Checklist of valid ant species known to occur in the Philippines, arranged alphabetically by genus. (Note: * denotes an unresolved homonym.)

There are 479 ant species known from the Philippines and at least an additional 100 undescribed or unidentified species represented by museum specimens and recent collections. The total size of the ant fauna is no doubt much larger, almost certainly more than 1000 species are present. This tentative list contains species names regarded as valid in Bolton 2013. It is meant to serve as a guide to our current knowledge of the Philippine ant fauna. A key to island codes is at the end of the list.

Species Distribution Notes

• Acanthomyrmex mindanao Moffett, 1986 Mnd
• Acropyga acutiventris Roger, 1862 Ceb, Mnd
• Acropyga nipponensis Terayama, 1985 Plw
• Acropyga pallida (Donisthorpe, 1938) Lzn A/G – new record
• Aenictus alticola Wheeler & Chapman (in Wheeler, 1930c) Lzn
• Aenictus appressipilosus Jaitrong & Yamane 2013 Palawan
• Aenictus bakeri Menozzi, 1925 Mnd
• Aenictus camposi Wheeler & Chapman, 1925 Lzn, Ngs
• Aenictus carolianus Zettel & Sorger, 2010b Ceb
• Aenictus ceylonicus (Mayr, 1866a) Lzn, Ngs
• Aenictus gracilis Emery, 1893b Lzn, Mnd, Mdr, Ngs
• Aenictus laeviceps F. Smith, 1857 Lyt, Lzn, Mnd, Ngs
• Aenictus luzoni Wheeler & Chapman, 1925 Lzn, Ngs
• Aenictus nesiotis Wheeler & Chapman (in Wheeler, 1930c) Lzn, Ngs, Mnd, Plw
• Aenictus pangantihoni Zettel & Sorger, 2010b Cam
• Aenictus philippinensis Chapman, 1963 Ngs
• Aenictus piercei Wheeler & Chapman (in Wheeler, 1930c) Mnd, Ngs
• Aenictus powersi Wheeler & Chapman (in Wheeler, 1930c) Ngs
• Aenictus rabori Chapman, 1963 Ngs
• Aenictus reyesi Chapman, 1963 Ngs
• Stigmatomma luzonicum Wheeler & Chapman, 1925 Lzn
• Stigmatomma reclinatum (Mayr, 1879) Lzn, Plw
• Stigmatomma rothneyi Forel, 1900c Lzn
• Anillomyrma decamera (Emery, 1901) Lzn
• Anochetus brevis Brown, 1978 Lzn, Mnd
• Anochetus cato Forel, 1901
• Anochetus graeffei Mayr, 1870 Lzn, Mnd, Ngs, Plw
• Anochetus incultus Brown, 1978 Lzn
• Anochetus isolatus Mann, 1919 Lzn, Ngs
• Anochetus leyticus Zettel, 2012 Lyt
• Anochetus modicus Brown, 1978 Lzn, Ngs
• Anochetus pangantihoni Zettel, 2012 Lyt
• Anochetus princeps Emery, 1884 Lzn
• Anochetus ruginotis Stitz, 1925 Lzn, Mnd
• Anochetus schoedli Zettel, 2012 Lzn
• Anochetus turneri Forel, 1900b Lzn
• Anochetus werneri Zettel, 2012 Mnd
• Anomalomyrma helenae Borowiec, Schulz, Alpert & Baňař, 2011 Plw
• Anoplolepis gracilipes (F. Smith, 1857) W
• Calyptomyrmex beccarii Emery, 1887b Lzn, Ngs
• Calyptomyrmex loweryi Shattuck, 2011 Lzn, Mnd
• Camponotus albocinctus (Ashmead, 1905) Lzn
• Camponotus barbosus Baroni Urbani, 1971 Lzn, Mnd
• Camponotus bellus Wheeler, 1919* Lzn
• Camponotus carin Emery, 1889 Lzn
• Camponotus castanicola Donisthorpe, 1943b Lzn, Mnd
• Camponotus cinerascens (Fabricius, 1787)
• Camponotus compressus (Fabricius, 1787) Plw
• Camponotus corallinus (Roger, 1863) Lzn, Mnd
• Camponotus gigas (Latreille, 1802) Lzn
• Camponotus horrens Forel, 1910 Lzn
• Camponotus irritans (F. Smith, 1857) Lzn
• Camponotus japonicus Mayr, 1866b Mnd
• Camponotus karawaiewi Menozzi, 1926 Mnd
• Camponotus leonardi Emery, 1889 Lzn, Ngs
• Camponotus maculatus Stitz, 1925* Lzn, Plw
• Camponotus nicobarensis Bingham, 1903* Lzn
• Camponotus nigricans Roger, 1863 Jol, Lzn
• Camponotus platypus Roger, 1863 Lzn, Mnd
• Camponotus pressipes Emery, 1893d Mnd
• Camponotus quadrisectus (F. Smith, 1858) Mnd, Plw
• Camponotus reticulatus Stitz, 1925* Lzn
• Camponotus rothneyi Viehmeyer, 1916b Lzn
• Camponotus rufifemur Stitz, 1925* Lzn
• Camponotus solenobius Menozzi, 1926 Mnd
• Camponotus trietericus Menozzi, 1926 Lzn
• Camponotus variegatus Forel, 1892a* Ngs
• Camponotus vitreus (F. Smith, 1860) Lzn
• Cardiocondyla kagutsuchi Terayama, 1999 Byg, Lzn
• Cardiocondyla mauritanica (Forel, 1890) Lzn
• Cardiocondyla sima Wheeler, 1935b Lzn, Mnd
• Cardiocondyla tjibodana Karavaiev, 1935 Byg, Lzn
• Cardiocondyla wroughtonii (Forel, 1890) Lzn
• Carebara alperti Fernandez, 2010 Ngs
• Cataulacus catuvolcus Bolton, 1974 Lzn, Rmb
• Cataulacus chapmani Bolton, 1974 Ngs
• Cataulacus setosus F. Smith, 1860b Mnd
• Centromyrmex feae (Emery, 1889) Bsn, Lzn
• Cerapachys bakeri (Wheeler & Chapman, 1925) Mnd
• Cerapchys biroi Forel, 1907
• Cerapachys bryanti Wheeler, 1919 Plw
• Cerapachys crawleyi Wheeler, 1924b Lzn, Ngs
• Cerapachys fragosus (Roger, 1862) Plw
• Cerapachys jacobsoni Forel, 1912a Ngs
• Cerapachys longitarsus (Mayr, 1879) Lzn
• Cerapachys luzuriagae (Wheeler & Chapman, 1925) Ngs
• Cerapachys muiri Wheeler & Chapman, 1925 Lzn
• Cerapachys pruinosus Brown, 1975 Ngs
• Cerapachys rufithorax Wheeler & Chapman, 1925 Lzn, Ngs
• Cerapachys sulcinodis Emery, 1889 Lzn, Ngs
• Cerapachys suscitatus (Viehmeyer, 1913) Ngs
• Crematogaster ampullaris F. Smith, 1861 Clg, Lyt, Lzn
• Crematogaster bakeri Menozzi, 1925 Mnd
• Crematogaster bicolor F. Smith, 1860b Lzn, Ngs
• Crematogaster brunnea Santschi, 1928
• Crematogaster crassicornis Emery, 1893d Lzn, Mnd
• Crematogaster cylindriceps Wheeler, 1927
• Crematogaster difformis F. Smith, 1857 Lzn
• Crematogaster inflata F. Smith, 1857 Lzn
• Crematogaster longiclava Emery, 1893d Lzn
• Crematogaster modiglianii Emery, 1900b Mnd, Ngs
• Crematogaster ochracea Mayr, 1862 Lzn, Ngs
• Crematogaster onusta Stitz, 1925 Plw, StC
• Crematogaster rogenhoferi Mayr, 1879 Lzn
• Crematogaster semperi Emery, 1893d Lzn
• Crematogaster simoni Emery, 1893d Lzn
• Crematogaster vitalisi Menozzi, 1925c Mnd
• Dacetinops cirrosus Taylor, 1985 Lyt HZW – new record
• Diacamma baguiense Wheeler & Chapman, 1925 Lzn
• Diacamma palawanicum Emery, 1900b Plw
• Diacamma panayense Wheeler & Chapman, 1925 Pny
• Diacamma rugosum (Le Guillou, 1842) W
• Diacamma sericeiventre Stitz, 1925 Plw
• Dilobocondyla chapmani Wheeler, 1924a Ngs
• Discothyrea bryanti (Wheeler, 1917) Lzn
• Discothyrea clavicornis Emery, 1897a Lzn
• Dolichoderus affinis Emery, 1889
• Dolichoderus patens Emery, 1900b Plw
• Dolichoderus thoracicus (F. Smith, 1860a) W
• Echinopla pallipes F. Smith, 1857 Lzn
• Echinopla pseudostriata (Donisthorpe, 1943a) Lzn
• Echinopla striata F. Smith, 1857
• Echinopla vermiculata Emery, 1898 Lzn
• Euprenolepis negrosensis (Wheeler, 1930b) Ngs
• Eurhopalothrix chapmani Taylor, 1990 Lzn
• Eurhopalothrix philippina Brown & Kempf, 1960 Lzn, Ngs
• Eurhopalothrix procera (Emery, 1897a) Lzn, Plw
• Forelophilus philippinensis Zettel & Zimmerman, 2007 Lzn, Mnd
• Forelophilus stefanschoedli Zettel & Zimmerman, 2007 Lyt, Lzn, Mnd
• Gauromyrmex acanthinus (Karavaiev, 1935) MCZ – new record
• Gesomyrmex luzonensis (Wheeler, 1916) Lzn, Ngs
• Gnamptogenys binghamii (Forel, 1900d) Lzn, Ngs, Plw
• Gnamptogenys bulbopila Lattke, 2004 Mnd
• Gnamptogenys chapmani Brown, 1958 Lzn, Ngs
• Gnamptogenys costata (Emery, 1889) Lzn, Mnd
• Gnamptogenys cribrata (Emery, 1900a) Lzn, Ngs
• Gnamptogenys fistulosa Lattke, 2004 Lzn
• Gnamptogenys laevior (Forel, 1905) Lzn
• Gnamptogenys leiolabia Lattke, 2004 Ngs
• Gnamptogenys luzonensis Wheeler, 1929d Lzn, Ngs
• Gnamptogenys menadensis (Mayr, 1887) Lzn
• Gnamptogenys posteropsis (Gregg, 1951) Lzn, Mnd, Ngs
• Harpegnathos empesoi Chapman, 1963 Mnd
• Harpegnathos macgregori Wheeler & Chapman, 1925 Bil
• Harpegnathos medioniger Donisthorpe, 1942 Lzn
• Harpegnathos venator Donisthorpe, 1937 Lzn
• Hypoponera confinis (Roger, 1860) Lzn
• Hypoponera opaciceps (Mayr, 1887)
• Hypoponera pruinosa (Emery, 1900a) Lzn, Ngs
• Hypoponera punctatissima (Roger, 1859) Lzn
• Hypoponera ragusai (Emery, 1894)
• Hypoponera sabronae (Donisthorpe, 1941b) Lzn
• Iridomyrmex anceps (Roger, 1863) Lzn, Mrq, Mnd
• Iridomyrmex angusticeps Forel, 1901 Mnd
• Lepisiota aurea Wheeler, 1935b Lzn, Ngs
• Lepisiota chapmani Wheeler, 1935b Plw
• Leptanilla astylina Petersen, 1968 Plw
• Leptogenys chinensis Mayr, 1870b
• Leptogenys diminuta (F. Smith, 1857) Lzn, Mnd, Ngs, Plw
• Leptogenys falcigera Roger, 1861 Ngs
• Leptogenys iridescens (F. Smith, 1857)
• Leptogenys maxillosa (F. Smith, 1858)
• Leptogenys mutabilis (F. Smith, 1861) Plw
• Leptogenys peuqueti Andre, 1887 Lzn, Ngs
• Leptogenys pruinosa Forel, 1900d
• Leptogenys punctiventris Mayr, 1879 Lzn
• Leptogenys varicosa Stitz, 1925 Plw
• Leptogenys watsoni Forel, 1900d
• Leptomyrmex fragilis (F. Smith, 1859) Mnd BPBM – new record
• Liomyrmex gestroi Emery, 1887b W
• Lophomyrmex bedoti Emery, 1893b Plw
• Lordomyrma diwata Taylor, 2012 Lzn
• Lordomyrma emarginata Taylor, 2012 Lzn
• Lordomyrma idianale Taylor, 2012 Lzn
• Lordomyrma limatula Taylor, 2012 Lyt
• Mayriella transfuga Baroni Urbani, 1977a Lzn A/G – new record
• Meranoplus biliran Schödl, 1998 Bil
• Metapone bakeri Wheeler, 1916 Lzn
• Metapone gracilis Wheeler, 1935b Mnd
• Monomorium banksi Forel, 1910 Ngs
• Monomorium destructor (Jerdon, 1851) Lzn, Plw
• Monomorium floricola Forel, 1910 Cam, Lzn, Mnd, Tbs
• Monomorium orientale Mayr, 1879
• Monomorium pharaonis (Linnaeus, 1758) W
• Myopias bidens Stitz, 1925 Ngs, Plw
• Myopias breviloba (Wheeler, 1919a) Bsn, Lzn, Ngs
• Myopias lobosa Willey & Brown, 1983 Lzn
• Myopias modiglianii (Emery 1900b) Ngs
• Myopias philippinensis Menozzi, 1925 Lzn, Mnd
• Myopopone castanea (F. Smith 1860b) Lyt, Lzn, Mnd, Ngs, Plw
• Myrmicaria aphidicola Calilung, 2000 Mnd
• Myrmicaria brunnea Saunders, 1842 W
• Myrmoteras glabrum Zettel & Sorger, 2011 Cam, Lzn
• Myrmoteras insulcatum Moffett, 1985 Lzn, Ngs
• Myrmoteras mcarthuri Zettel & Sorger, 2011 Lyt, Lzn
• Myrmoteras williamsi Wheeler, 1919a Lzn, Ngs
• Mystrium camillae Emery, 1889 Lzn, Plw
• Noonilla copiosa Petersen, 1968 Plw
• Ochetellus glaber (Mayr, 1862) Lzn, Ngs MCZ – new record
• Odontomachus alius Sorger & Zettel, 2011 Bhl, Bil, Ceb, Cat, Lyt, Smr
• Odontomachus banksi Forel, 1910 Ceb, Lzn
• Odontomachus infandus F. Smith, 1858 Lzn, Mdr
• Odontomachus malignus F. Smith, 1859 Ban, Bhl, Jol, Twi
• Odontomachus papuanus Emery, 1887b Ngs, Plw, Pny
• Odontomachus philippinus Emery, 1893d Ngs, Pny, Sqr
• Odontomachus rixosus F. Smith, 1859 Mnd, Plw
• Odontomachus schoedli Sorger & Zettel, 2011 Lzn
• Odontomachus scifictus Sorger & Zettel, 2011 Cam
• Odontomachus simillimus F. Smith, 1858 W
• Odontoponera denticulata (F. Smith, 1858) W
• Oecophylla smaragdina (Fabricius, 1775) W
• Overbeckia subclavata Viehmeyer, 1916a MCZ – new record
• Pachycondyla annamita (Andre, 1892) Lzn
• Pachycondyla chinensis (Emery, 1895b)
• Pachycondyla claudata (Menozzi, 1926) Lzn, Smr
• Pachycondyla croceicornis (Emery, 1900a)
• Pachycondyla darwinii (Emery, 1899) Lzn, Ngs
• Pachycondyla glabripes (Emery, 1893d) Mnd
• Pachycondyla javana (Mayr 1867)
• Pachycondyla leeuwenhoeki (Forel, 1886a) Ngs MCZ – new record
• Pachycondyla luteipes (Mayr, 1862) Lzn
• Pachycondyla melancholica F. Smith, 1865
• Pachycondyla myropola (Menozzi, 1925) Lzn
• Pachycondyla obscurans (Walker, 1859) Lzn
• Pachycondyla stigma (Fabricius, 1804)
• Pachycondyla testacea (Emery, 1893a) Lzn A/G – new record
• Pachycondyla tridentata F. Smith, 1858 Jol, Twi
• Pachycondyla williamsi (Wheeler & Chapman, 1925) Lzn
• Paraparatrechina iridescens (Donisthorpe, 1942) Lzn, Plw
• Paratopula ceylonica (Emery, 1901)
• Paratopula macta Bolton, 1988 Lzn
• Paratopula sumatrensis (Forel, 1913)
• Paratrechina longicornis (Latreille, 1802) W
• Pheidole aglae Forel, 1913 Lzn
• Pheidole aristotelis Forel, 1911a Ngs, Plw
• Pheidole cariniceps Eguchi, 2001a Lzn A/G – new record
• Pheidole clypeocornis Eguchi, 2001a Lzn
• Pheidole elisae Emery, 1900 Mnd
• Pheidole fantasia Chapman, 1963 Lzn, Ngs
• Pheidole fervens F. Smith, 1858 Lzn, Ngs
• Pheidole hortensis Forel, 1913 Lzn, Smr
• Pheidole inscrobiculata Viehmeyer, 1916a Lzn
• Pheidole jacobsoni Forel, 1911b Lzn, Mnd, Smr
• Pheidole kikutai Eguchi, 2001 Mnd
• Pheidole lokitae Forel, 1913 Mnd
• Pheidole longipes (F. Smith, 1857) Mnd
• Pheidole maculifrons Wheeler, 1929d Lzn
• Pheidole makilingi Viehmeyer, 1916b Lzn
• Pheidole megacephala (Fabricius, 1793)
• Pheidole parva Mayr, 1865 Mnd
• Pheidole plagiaria Stitz, 1925 Lyt, Plw
• Pheidole quadricuspis Emery, 1900a Lzn
• Pheidole sarawakana Forel, 1911a Lzn
• Pheidole sauberi Forel, 1905 Lzn
• Pheidole sayapensis Eguchi, 2001a Lzn
• Pheidole simoni Emery, 1893d Lzn
• Pheidole singaporensis Özdikmen, 2010 Mnd
• Pheidole tjibodana Forel, 1905 Lzn
• Pheidologeton affinis (Jerdon, 1851) Plw
• Pheidologeton diversus (Jerdon, 1851) Lzn
• Pheidologeton maccus Wheeler, 1929d Lzn, Smr
• Pheidologeton pygmaeus Emery, 1893d Lzn
• Pheidologeton silvestrii Wheeler, 1929d Lzn
• Philidris myrmecodiae (Emery, 1887a) Lzn
• Platythyrea bidentata Brown, 1975 Ngs
• Platythyrea inermis Forel, 1910 Lzn, Ngs, Smr
• Platythyrea parallela (F. Smith, 1859) Lzn, Mnd, Ngs, Rmb
• Platythyrea quadridenta Donisthorpe, 1941 Plw
• Polyrhachis abdominalis F. Smith, 1858 Pl
• Polyrhachis aculeata Mayr, 1879 Ng
• Polyrhachis aequalis Forel, 1910 Lz, Ng, Pl
• Polyrhachis arcuata (Le Guillou, 1842)
• Polyrhachis armata (Le Guillou, 1842) Lz, Md, Ng, Pl
• Polyrhachis baca Sorger & Zettel, 2010 Lz
• Polyrhachis bakeri Viehmeyer, 1916b Lz
• Polyrhachis banghaasi Viehmeyer, 1922
• Polyrhachis bellicosa F. Smith, 1859 Lz, Ng
• Polyrhachis bicolor F. Smith, 1858 Lz, Pl
• Polyrhachis bihamata (Drury, 1773) Lz, Md, Ng
• Polyrhachis carbonaria F. Smith, 1857
• Polyrhachis chapmani Kohout, 2006b Ng
• Polyrhachis creusa Emery, 1897b Ng
• Polyrhachis cryptoceroides Emery, 1887a Lz
• Polyrhachis cyaniventris F. Smith, 1858 Lz
• Polyrhachis diana Wheeler, 1909 Md
• Polyrhachis dives F. Smith, 1857 Lz, Pl
• Polyrhachis empesoi Kohout, 2006b Ng
• Polyrhachis esuriens Emery, 1897b Ng
• Polyrhachis etheli Chapman, 1963 Ng
• Polyrhachis exotica Kohout, 1987 Md
• Polyrhachis follicula Menozzi, 1926 Lz
• Polyrhachis hector F. Smith, 1857 Ng
• Polyrhachis hippomanes Stitz, 1925* Lz, Md, Ng
• Polyrhachis ignota Kohout, 1987 Lz
• Polyrhachis illaudata Walker, 1859 Lz, Pl
• Polyrhachis inermis F. Smith, 1858 Ng, Pl
• Polyrhachis javanica Mayr, 1867
• Polyrhachis lama Kohout, 1994b Lz
• Polyrhachis magnifica Menozzi, 1926 Lz, Ms, Ng, Tb
• Polyrhachis marginata F. Smith, 1859
• Polyrhachis mindanaensis Emery, 1923 Lz, Md
• Polyrhachis mitrata Menozzi, 1932 Md
• Polyrhachis mucronata F. Smith, 1859
• Polyrhachis muelleri Forel, 1893a Md, Ng
• Polyrhachis murina Emery, 1893b Lz, Sm
• Polyrhachis nigropilosa Mayr, 1872 Ng, Pl
• Polyrhachis noesaensis Forel, 1915 Ng, Pl, Si
• Polyrhachis obesior Viehmeyer, 1916a
• Polyrhachis oedocantha Wheeler, 1919a Ng
• Polyrhachis olybria Forel, 1912b Lz. Ng, Md
• Polyrhachis orpheus Forel, 1911b
• Polyrhachis osiris Bolton, 1975 Lz, Ng
• Polyrhachis parabiotica Chapman, 1963 Lz, Ng
• Polyrhachis pellita Menozzi, 1922 Lz, Ng, Sm
• Polyrhachis phalerata Menozzi, 1926 Lz
• Polyrhachis philippinensis F. Smith, 1858 Lz
• Polyrhachis pirata Sorger and Zettel, 2009 Lz, Mr, Sm
• Polyrhachis pressa Mayr, 1862 Md
• Polyrhachis pubescens Mayr, 1879 Lz
• Polyrhachis rufipes F. Smith, 1858 Lz
• Polyrhachis saevissima F. Smith, 1860a Lz, Md, Ng
• Polyrhachis saigonensis Forel, 1886b Lz
• Polyrhachis scabra Kohout, 1987 Md
• Polyrhachis sculpturata F. Smith, 1860a Lz, Pl
• Polyrhachis semiinermis Donisthorpe, 1941c Lz, Md
• Polyrhachis sexspinosa (Latreille, 1802) Sm
• Polyrhachis solivaga Menozzi, 1926 Lz
• Polyrhachis striata Mayr, 1862 Pl
• Polyrhachis thrinax Roger, 1863 Lz
• Polyrhachis tragos Stitz, 1925 Lz, Md
• Polyrhachis tubifex Karavaiev, 1926 Lz
• Polyrhachis venus Forel, 1893 Lz, Md
• Polyrhachis villipes F. Smith, 1857 Md
• Polyrhachis vindex F. Smith, 1857 Lz, Ng
• Polyrhachis ypsilon Emery, 1887a Lz, Ng
• Polyrhachis zopyra F. Smith, 1861 Lz, Ng
• Ponera chapmani Taylor, 1967 Lz, Md, Sm
• Ponera oreas (Wheeler, 1933) Ng
• Prionopelta kraepelini Forel, 1905 Lz, Ng, Sm UPLB
• Pristomyrmex bicolor Emery, 1900b Lz, Pl
• Pristomyrmex brevispinosus Emery, 1887b Md
• Pristomyrmex cebuensis Zettel, 2007 Cb
• Pristomyrmex collinus Wang, 2003 Lz, Md, Ng, Pn, Tb
• Pristomyrmex costatus Wang, 2003 Md
• Pristomyrmex curvulus Wang, 2003 Ng
• Pristomyrmex distinguendus Zettel, 2006 Ly, Lz
• Pristomyrmex divisus Wang, 2003 Ng
• Pristomyrmex flatus Wang, 2003 Lz
• Pristomyrmex hirsutus Wang, 2003 Md
• Pristomyrmex levigatus Emery, 1897a Lz
• Pristomyrmex longispinus Wang, 2003 Ng
• Pristomyrmex picteti Emery, 1893d Ly, Lz, Md, Ng, Pl
• Pristomyrmex punctatus (F. Smith, 1860b) Ly, Md, Mq, Mr, Pc, Sm
• Pristomyrmex quadridens Emery, 1897a Ly
• Pristomyrmex rugosus Zettel, 2006 Ly
• Pristomyrmex schoedli Zettel, 2006 Ly
• Pristomyrmex simplex Wang, 2003 Lz
• Probolomyrmex dammermani Wheeler, 1928 Ng MCZ – new record
• Proceratium papuanum Emery, 1897a Lz A/G – new record
• Pseudolasius typhlops Wheeler, 1935b Lz, Pl
• Pyramica dohertyi (Emery, 1897a) Lz
• Pyramica jacobsoni (Menozzi, 1939) Pl
• Pyramica karawajewi (Brown, 1948) Lz
• Pyramica mitis Brown, 2000 Lz
• Pyramica pedunculata (Brown, 1953) Lz, Ng
• Pyramica scylla Bolton, 2000 Lz
• Pyramica serradens Bolton, 2000 Lz
• Recurvidris nigrans Zettel, 2008 Ng
• Rhoptromyrmex wroughtonii Forel, 1902a Lz MCZ, A/G
• Rhytidoponera araneoides (Le Guillou, 1842) Ng BPBM– new record
• Romblonella opaca (F. Smith, 1861) Pl, Rr, Rm
• Simopone chapmani Taylor, 1966 Ng
• Solenopsis geminata (Fabricius, 1804) W
• Strumigenys arrogantia Bolton, 2000 Lz, Pl
• Strumigenys chapmani Brown, 1954 Lz, Ng
• Strumigenys emmae (Emery, 1890b) Lz
• Strumigenys esrossi Brown, 1957 Lz
• Strumigenys godeffroyi Mayr, 1866a Lz, Md, Pl
• Strumigenys imantodes Bolton, 2000 Lz
• Strumigenys indigatrix Wheeler, 1919a Pl
• Strumigenys inhonesta Bolton, 2000 Lz
• Strumigenys juliae Forel, 1905 Pl
• Strumigenys koningsbergeri Forel, 1905 Lz, Pl
• Strumigenys lewisi Cameron, 1886 Lz A/G – new record
• Strumigenys mirifica Bolton, 2000 Lz
• Strumigenys morphica Bolton, 2000 Lz A/G – new record
• Strumigenys mododonta Bolton, 2000 Lz
• Strumigenys perplexa (F. Smith, 1876) Lz A/G – new record
• Strumigenys phytibia Brown, 1957
• Strumigenys rantan Bolton, 2000 Md
• Strumigenys rogeri Emery, 1890b
• Strumigenys signeae Forel, 1905 Pl
• Strumigenys synchysis Bolton, 2000 Lz A/G – new record
• Strumigenys szalayi Emery, 1897a Lz, Md, Mr, Ng
• Strumigenys tenitecta Bolton, 2000 Lz
• Strumigenys uichancoi Brown, 1957
• Tapinoma emeryi (Ashmead, 1905) Lz SI
• Tapinoma melanocephalum (Fabricius, 1793) W
• Tapinoma philippinense Donisthorpe, 1942 Lz
• Tapinoma williamsi (Wheeler, 1935b) Lz
• Technomyrmex albipes (F. Smith, 1861) Lz, Mr
• Technomyrmex difficilis Forel, 1892b By, Lz, Md
• Technomyrmex elatior Forel, 1902b Lz, Md
• Technomyrmex grandis Emery, 1887a Bh, Ly, Md, Ng
• Technomyrmex kraepelini Forel, 1905 Lz
• Technomyrmex pratensis (F. Smith, 1860b) Md
• Technomyrmex sundaicus (Emery, 1900b) Lz, Md, Sy
• Technomyrmex textor Forel, 1909 Lz, Pl
• Technomyrmex vitiensis Mann, 1921 Pl
• Technomyrmex wheeleri (Emery, 1913a) Ly, Lz, Rm
• Tetramorium aspersum (F. Smith, 1865) Lz, Md, Ng
• Tetramorium bicarinatum (Nylander, 1846) W
• Tetramorium chapmani Bolton, 1977 Ng
• Tetramorium cynicum Bolton, 1977 Ng
• Tetramorium eleates Forel, 1913 Lz
• Tetramorium guineense (Bernard, 1953) Lz, Ng, Pl, Sm
• Tetramorium insolens (F. Smith, 1861) Ng
• Tetramorium katypum (Bolton, 1976) Cm, Lz, Md
• Tetramorium kheperra (Bolton, 1976) Md
• Tetramorium khnum Bolton, 1977 Ng
• Tetramorium kraepelini Forel, 1905 Ng
• Tetramorium lanuginosum Mayr, 1870 Ng
• Tetramorium laparum Bolton, 1977 Ng
• Tetramorium manobo (Calilung, 2000) Md
• Tetramorium obtusidens Viehmeyer, 1916a
• Tetramorium pacificum Mayr, 1870 Ly, Lz, Md, Ng, Rm
• Tetramorium parvispinum (Emery, 1893b) Lz
• Tetramorium rinatum Bolton, 1977 Ng
• Tetramorium simillimum (F. Smith, 1851) Ng
• Tetramorium smithi Mayr, 1879
• Tetramorium tonganum Mayr, 1870
• Tetramorium tortuosum Roger, 1863
• Tetramorium walshi Forel 1890
• Tetramorium zypidum Bolton, 1977 Ng
• Tetraponera allaborans (Walker, 1859) W
• Tetraponera attenuata F. Smith, 1877 Pl
• Tetraponera difficilis (Emery, 1900b) W
• Tetraponera extenuata Ward, 2001 Lz, Md
• Tetraponera inversinodis Ward, 2001 Pl
• Tetraponera modesta (F. Smith, 1860b) Ng, Pl, Rm, Sy, Tb
• Tetraponera nigra (Jerdon, 1851) Pl
• Tetraponera nitida (F. Smith, 1860b) W
• Tetraponera pilosa (F. Smith, 1858) Pl
• Vollenhovia ambitiosa Menozzi, 1925 Lz, Md
• Vollenhovia banksi Forel, 1910 Ng
• Vollenhovia cristata (Stitz, 1938) Pl
• Vollenhovia luctuosa (Stitz, 1938) Pl
• Vollenhovia oblonga Forel, 1910 Lz
• Vollenhovia soleaferrea Donisthorpe, 1942 Lz
• Vombisidris philippina Zettel & Sorger, 2010a Cb

Key to Notes:

• A/G = Alpert and General Collection
• BPBM = Bernice P. Bishop Museum Collection
• HZW = Herbert Zettel (Vienna) Collection
• MCZC = Museum of Comparative Zoology Collection
• UPLB = University of the Philippines at Los Baños Museum of Natural History Collection
• SI = Smithsonian Institution Museum Collection

Key to Island Distribution:

• Ban = Bantayan
• Bbc = Balabac
• BcG = Bucas Grande
• Bhl = Bohol
• Bil = Biliran
• Bsn = Basilan
• Bus = Busuanga
• Byg = Bayagnan
• Ceb = Cebu
• Clg = Calabugtong
• Cam = Camiguin
• Cat = Catanduanes
• Din = Dinagat
• Hik = Hikdop
• Jol = Jolo
• Lyt = Leyte
• Lzn = Luzon
• Mnd = Mindanao
• Mrq = Marinduque
• Mdr = Mindoro
• Msb = Masbate
• Ngs = Negros
• Pac = Pacijan
• Plw = Palawan
• Pny = Panay
• Pol = Polillo
• Rmb = Romblon
• Rrp = Rapu-rapu
• StC = Sta. Cruz
• Sbt = Sibutu
• Smr = Samar
• Sqr = Siquijor
• Sby = Sibuyan
• Tbs = Tablas
• Twi = Tawi-tawi
• W = Widespread