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

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
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 Taxonomic references are frequently hard to access, badly outdated, and difficult for the non-specialist to use.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.

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

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., 1 known from PH.

Only 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). 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).

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

Genus Myrmoteras Forel, 1893 – Formicinae: 32 spp., at least 4 known from PH. 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), [[Media:Zettel & Sorger 2011.pdf|Zettel and Sorger (2011)]] Philippine species.
 * Myrmoteras glabrum
 * Myrmoteras insulcatum
 * Myrmoteras mcarthuri
 * Myrmoteras williamsi

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).

Noonilla
Genus Noonilla Petersen, 1968 – Leptanillinae: 1 sp., 2 known from PH.

Described from an alate male reproductive, collected in southern Palawan Island, Noonilla copiosa Petersen, 1968 is the only described species of this genus. 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 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).

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
31 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.

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