Tapinoma melanocephalum

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Tapinoma melanocephalum
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Dolichoderinae
Genus: Tapinoma
Species: T. melanocephalum
Binomial name
Tapinoma melanocephalum
(Fabricius, 1793)

Tapinoma melanocephalum casent0173215 profile 1.jpg

Tapinoma melanocephalum casent0173215 dorsal 1.jpg

Specimen Label


Common Name
Ghost Ant
Language: English
Language: Japanese

This is a widespread tropical tramp species, introduced throughout the world. Nests are found in the soil, under stones, fallen trees, tree bark and in other crevices. Its native range is unknown. It is ubiquitous in non-air conditioned dwellings anywhere in the lowland tropics. Regardless where you are, if you are sitting at a table with a sugar dispenser you are likely to see workers of T. melanocephalum running about on the surface. They always seem to be able to find their way into the sugar container, and sugar on tropical tables always contains some non-negligible fraction of T. melanocephalum workers. When you put a spoonful of sugar in your drink, you can judge the level of contamination by how many workers are left floating on the surface.

In quantitative biodiversity surveys, this species often has to be excluded from data analysis because the laboratory where samples are processed contains T. melanocephalum as a pest, and contamination of samples occurs.

Although most often found in houses, they can also move out into surrounding vegetation in highly disturbed and highly insolated habitats, opportunistically nesting in small plant cavities. Nests readily relocate, and overnight they can move into a shoe or an umbrella left on a porch. (Jack Longino)

At a Glance • Highly invasive  • Polygynous  • Supercolonies  

Photo Gallery

  • Tapinoma melanocephalum workers recruited to a food source (Kerala, India). Photo by Kalesh Sadasivan.


Distinctively bicoloured, head brown, alitrunk and gaster pale yellowish with variable brown patches; legs and antennae very pale. Prothorax laterally compressed, broadening anteriorly. Length: 1.5-2 mm. (Collingwood 1979)

Keys including this Species


Tapinoma melanocephalum has one of the widest distributions known for any ant species. It has spread across the Old World and New World in both the northern and southern hemispheres. It is tropical and subtropical; at latitudes greater than 30°, it is largely restricted to living inside buildings. The only ants with broader known distributions than T. melanocephalum are the longhorn crazy ant, Paratrechina longicornis, and the pharaoh ant, Monomorium pharaonis. I expect that T. melanocephalum will eventually spread to the disturbed lowlands of virtually every tropical country. This species also appears to be an increasingly common indoor pest in temperate areas, and some-day may be found in every major city of the world. Because T. melanocephalum can live anywhere that humans live, there is no limit to the latitude where indoor populations of this species may be found. (Wetterer 2009)

Latitudinal Distribution Pattern

Latitudinal Range: 32.812778° to -9.233°.

Tropical South

Distribution based on Regional Taxon Lists

Afrotropical Region: Benin, Cape Verde, Comoros, Gambia, Guinea, Nigeria, Saint Helena, Senegal, Socotra Archipelago, United Arab Emirates, Yemen.
Australasian Region: Australia, New Caledonia, Norfolk Island.
Indo-Australian Region: Borneo, Cook Islands, Fiji, French Polynesia, Guam, Hawaii, Indonesia (type locality), Krakatau Islands, Malaysia, New Guinea, Niue, Philippines, Samoa, Singapore, Solomon Islands, Tokelau, Tonga, Vanuatu (type locality), Wallis and Futuna Islands.
Malagasy Region: Madagascar, Mauritius, Mayotte, Réunion, Seychelles.
Nearctic Region: Canada, United States.
Neotropical Region: Anguilla, Bahamas, Barbados, Belize, Brazil, Cayman Islands, Colombia (type locality), Costa Rica (type locality), Cuba, Dominican Republic, Ecuador, French Guiana, Galapagos Islands, Greater Antilles, Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Lesser Antilles, Mexico, Netherlands Antilles, Paraguay, Puerto Rico, Suriname, Trinidad and Tobago.
Oriental Region: Bangladesh, Cambodia, India (type locality), Laos, Maldives, Nicobar Island, Pakistan, Sri Lanka, Thailand.
Palaearctic Region: Afghanistan, Belgium, Canary Islands, China, Czech Republic, Finland, Hungary, Iberian Peninsula, Iceland, Italy, Japan, Oman, Republic of Korea, Romania, Russian Federation, Spain, Ukraine, United Kingdom of Great Britain and Northern Ireland.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.


San Cristóbal, República Dominicana. Video by Judá Isaí Martínez Uribe.

Wetterer (2009) - "Tapinoma melanocephalum is a ubiquitous pest through much of the tropics and subtropics. As Longino (2006) aptly wrote: "regardless of whether you are in Guinea, New Guinea, or Guyana, if you are sitting at a table with a sugar dispenser you are likely to see workers of T. melanocephalum running about on the surface." Workers are small (~ 1.5 mm) and their pale legs and abdomens often blend into the background, making them difficult to see. Their dark brown heads and thoraces often look like hovering specks, unrecognizable as ants. The barely visible "ghostly" appearance of T. melanocephalum no doubt explains its common name.

Tapinoma melanocephalum is particularly noted as a pest in homes, restaurants, hospitals, and greenhouses. For example, T. melanocephalum was the most common house ant in Brazil (e.g., Kamura et al. 2007) and on Panang Island, Malaysia (Lee et al. 2002), and it was the most fre-quently reported pest ant in greater Miami (Klotz et al. 1995). Wheeler (1907) noted that T. melanocephalum had been "introduced into well-heated buildings in New York," and observed workers in the winter "visiting the sugar bowls on the tables of the Columbia University commons." Karawajew (1926) reported T. melanocephalum from hotel tables both on Ambon Island, Indonesia and in Vladivostok, Russia. James Trager (pers. comm.) observed T. melanocephalum inside the Sachs Butterfly House in Chesterfield, Missouri, beginning shortly after it opened in 1995. Tapinoma melanocephalum also has been a long term problem in the greenhouses of the Dow Estate in Midland, Michigan, where fumigation is incompatible with a butterfly exhibit (J. LaForest, pers. comm.).

Although T. melanocephalum does not sting and its bite is not very painful (pers. obs.), it can be a serious pest. For example, Fowler et al. (1993) and Moreira et al. (2005) found that T. melanocephalum was the most prevalent ant in hospitals in Brazil, and had the highest potential as a vector for pathogens. In fact, Moreira et al. (2005) found at least 14 different types of bacteria on T. melanocephalum workers collected in the hospitals, including antibiotic resistant strains. Tapinoma melanocephalum also can be an agricultural pest because it tends populations of phloem feeding Hemiptera, such as mealybugs, scale insects, and aphids, for their honeydew (Venkataramaiah and Rehman 1989). Hemiptera cause damage by sapping plants of nutrients and increasing the occurrence of diseases, including viral and fungal infections. Fowler et al. (1994) concluded that T. melanocephalum had a great impact on other ant species on banana plantations in Brazil. Under some conditions, T. melanocephalum may be beneficial as a biocontrol agent. For example, Osborne et al. (1995) found that in greenhouses of central Florida, naturally occurring populations of T. melanocephalum were important in controlling a serious plant pest, the two-spotted spider mite, Tetranychus urticae Koch, 1836.

Whereas T. melanocephalum is most prevalent in disturbed habitats, it has also been reported from natural reserves around the world, including the Baiyer River Wildlife Sanctuary in Papua-New Guinea (Whalen and Mackay 1988), Reserva Biológica de Una in Brazil (Delabie et al. 1997), primary rainforest in Chocó, Colombia (Armbrecht et al. 2001), and on many Galapagos islands (Causten et al. 2006). In the Wapoga River Area of Irian Jaya, Indonesia, Snelling (2000) found that T. melanocephalum "had successfully invaded forest areas where they apparently became quickly assimilated into the native mix; their presence did not seem to adversely affect other ant species there. This apparently benign situation probably would be reversed in the presence of continued disturbance of the habitat."

Tapinoma melanocephalum colonies are polygynous and frequently relocate their nests. In a matter of hours, colonies can move into cargo, which is then shipped worldwide. In fact, Lester (2005) reported that T. melanocephalum was the second most common ant species (after Pheidole megacephala) intercepted in freight entering New Zealand between 1997 and 2002. Weber (1939) reported collecting T. melanocephalum from riverboats in Venezuela and Guyana and on a ship between Haiti and New York. Tapinoma melanocephalum was found on a train in France, traveling from Paris to Rennes (X. Espadaler, pers. comm.). In Le Carbet, Martinique, a T. melanocephalum colony moved overnight into luggage packed for departure, requiring me to shake out everything before repacking."

Klimes and Okrouhlik (2015) - One of the most frequently recorded pests in hospitals in the tropics . . . In buildings, it is usually seen in kitchen corridors and bathrooms . . . Despite their diminutive size, workers are able to use their mandibles to penetrate even the thick plastic of sugar packages.

Although this species has been listed as supercolonial (Debout et al., 2007; Eyer & Vargo, 2021), genetic evidence of this is lacking.

Regional Notes

Espadaler (2007) - Canary Islands: Running workers were detected in irrigated gardens next to recently built bungalows. This is the first record for El Hierro of this well known tramp species.

Klimes and Okrouhlik (2015) report on a indoor infestation in an apartment complex in South Bohemia, Czech Republic. This is the first time Tapinoma melanocephalum has been recorded from the country.

This species is common in south Florida, to Volusia county. Almost always in disturbed areas, in many sites seems confined to areas around buildings. Nests are in bark mulch, under objects on the ground, and under loose bark and the bases of palm fronds. Pest status: can enter buildings through screens and small cracks, becoming a general annoyance. First published Florida record: Smith 1930. (Deyrup, Davis & Cover, 2000.)

Sharaf et al. (2017) - Yemen: The nesting and foraging habitats of this species are diverse. Several specimens were found foraging on a tree located on a mountainside with nearby stream drainage. The area had moist soils with high plant diversity, especially Adiantum capillus-veneris L. (Pteridaceae), which was abundant. The area was dominated by the ponerine ant, Brachyponera sennaarensis (Mayr, 1862). A nest series was collected from an area with moist soil and dense grasses. The area was rich in decayed animal faeces. Another nest was collected by sifting the leaf litter next to a stream and small pool with many scattered date palm (Phoenix dactylifera L.) trees. Another nest series was found under the bark of a recently cut dragon blood tree, Dracaena cinnabari Balf.f. (Asparagaceae) and associated with curculionid beetles. This invasive species has a broad range of habitat preference worldwide (Wetterer 2009a), including pre-existing cavities made of plant materials, and in larders and pantries, and nesting in walls and potted plants indoors (Ellison et al. 2012).


An experimental study by Zhou et al. (2015) tested effects of ant tending, host plants, parasitism and their interactions on mealybugs on two host plant species. The plants, tomato (Solanum lycopersicun) and cotton (Gossypium hirsutum), were grown in a greenhouse at an agricultural field station near Guangzhou, China. The ants (Tapinoma melanocephalum), hemipterans (Phenacoccus solenopsis Tinsley) and parasitoid (Aenasius bambawalei Hayat) were collected from local habitats, i.e., are naturally sourced and locally occurring species that are known to interact with one another. They found that mealybug benefits from ant attendance by T. melanocephalum did differ by host plant. The ant tending was more intense on tomato versus cotton plants. The mealybugs showed both a higher growth rate with higher ant attendance and a decrease in parasitism rate. In sum, they found the tomato plants likely produced higher quality honeydew, stimulating ant attendance, and this in turn provided positive benefits to the hemiptera.

Cheng et al. (2019), studying this species in Guangzhou China, found a number of differences in workers sampled from populations of T. melanocephalum from areas with Solenopsis invicta versus areas where this other ant is not present. For many analyses and experiments that were part of this study, Tapinoma melanocephalum workers were sampled from numerous colonies within each of three sites with S. invicta and three sites without. It appears competition between these two invasive species alters T. melancephalum aggressiveness (less aggressive when S. invicta is present), diet (stable isotype analysis suggests a switch from a carbohydrate-rich to a protein-rich diet when S. invicta is present), microbial bacteria symbionts, and other bacteria (the relative abundance of Wolbachia, Lactobacillales and Enterobacteriaceae). Being less aggressive potentially reduces harmful interactions between T melanocephalum and S. invicta. The change in diet, and a shift away from nectar and honeydew sources that are likely exploited by S. invicta, is an apparent adjustment to avoid competition (or is simply a result of competing poorly). This dietary change in turn putatively alters the workers microbial symbionts and bacteria.

Association with Other Organisms

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Other Insects

This ant has been observed tending larvae of Lampides boeticus (Obregon et al. 2015).

This species is known to tend the following aphids in Pakistan (Siddiqui et al., 2019; Gull-E-Fareen et al., 2020): Aphis eugeniae, Aphis fabae, Aphis fabae solanella, Aphis gossypii, Aphis pomi, Greenidea (Greenidea) decaspermi, Greenidea (Greenidea) ficicola, Greenidea (Trichosiphum) formosana, Hysteroneura setariae, Macrosiphoniella sanborni, Myzus persicae, Ovatus crataegarius, Pentalonia nigronervosa, Tinocallis kahawaluokalani and Tuberolachnus salignus.


Shi et al. (2017) - Abstract Volatile compounds in Tapinoma melanocephalum (Fabricius) (Hymenoptera: Formicidae, Dolichoderinae) workers were analyzed using gas chromatography–mass spectrometry and electroantennogram responses of workers to these compounds were investigated using coupled gas chromatography–electroantennographic detection. Among 18 detected compounds, six elicited electroantennogram response, including 6-methyl-5-hepten-2-one, 4 iridodials, and (Z)-9-nonadecene. (Z)-9-Nonadecene and iridodials were identified the first time in T. melanocephalum. (Z)-9-Nonadecene is a major component in mandibular glands and iridodials are produced in pygidial glands. In contrast to previous report, actinidine was not found in pygidial glands. Behavioral response of workers to the synthetic (Z)-9-nonadecene was investigated. (Z)-9-nonadecene is an attractant to T. melanocephalum. It also affected their locomotion patterns.

Life History Traits

  • Queen number: polygynous (Holldobler & Wilson, 1977)



MCZ-ENT00512185 Tapinoma melanocephalum melanocephalum hef.jpgMCZ-ENT00512185 Tapinoma melanocephalum melanocephalum hal 5.jpgMCZ-ENT00512185 Tapinoma melanocephalum melanocephalum had 5.jpgMCZ-ENT00512185 Tapinoma melanocephalum melanocephalum lbs.jpgTapinoma melanocephalum casent0008659 head 1.jpgTapinoma melanocephalum casent0008659 profile 1.jpgTapinoma melanocephalum casent0008659 dorsal 1.jpgTapinoma melanocephalum casent0008659 label 1.jpgTapinoma melanocephalum casent0125363 head 1.jpgTapinoma melanocephalum casent0125363 profile 1.jpgTapinoma melanocephalum casent0125363 dorsal 1.jpgTapinoma melanocephalum casent0125363 label 1.jpgTapinoma melanocephalum casent0005325 head 1.jpgTapinoma melanocephalum casent0005325 profile 1.jpgTapinoma melanocephalum casent0005325 dorsal 1.jpgTapinoma melanocephalum casent0005325 label 1.jpgTapinoma melanocephalum casent0173744 head 1.jpgTapinoma melanocephalum casent0173744 profile 1.jpgTapinoma melanocephalum casent0173744 dorsal 1.jpgTapinoma melanocephalum casent0173744 label 1.jpg

Additional worker images can be found here


Images from AntWeb

Tapinoma melanocephalum casent0444732 head 1.jpgTapinoma melanocephalum casent0444732 profile 1.jpgTapinoma melanocephalum casent0444732 dorsal 1.jpgTapinoma melanocephalum casent0444732 label 1.jpg
Queen (alate/dealate). Specimen code casent0444732. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Tapinoma melanocephalum casent0173214 head 1.jpgTapinoma melanocephalum casent0173214 profile 1.jpgTapinoma melanocephalum casent0173214 dorsal 1.jpgTapinoma melanocephalum casent0173214 label 1.jpg
Queen (alate/dealate). Specimen code casent0173214. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CDRS, Galapagos, Ecuador.
Tapinoma melanocephalum casent0125327 head 1.jpgTapinoma melanocephalum casent0125327 profile 1.jpgTapinoma melanocephalum casent0125327 dorsal 1.jpgTapinoma melanocephalum casent0125327 label 1.jpg
Queen (alate/dealate). Specimen code casent0125327. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


Tapinoma melanocephalum iccdrs0002606 head 1.jpgTapinoma melanocephalum iccdrs0002606 profile 1.jpgTapinoma melanocephalum iccdrs0002606 profile 2.jpgTapinoma melanocephalum iccdrs0002606 dorsal 1.jpgTapinoma melanocephalum iccdrs0002606 label 1.jpg


The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • melanocephalum. Formica melanocephala Fabricius, 1793: 353 (w.) FRENCH GUIANA.
    • [Misspelled as melanorephalum by Santschi, 1924c: 112.]
    • Emery, 1887a: 249 (m.); Forel, 1891b: 102 (q.); Wheeler, G.C. & Wheeler, J. 1951: 197 (l.); Crozier, 1970: 119 (k.).
    • [Note: neotype w. designated by Guerrero, 2018: 499. Colombia: Magdalena. Guerrero, 2018: 499, also nominates “paraneotypes”: under ICZN rules no such category exists.]
    • Combination in Lasius: Fabricius, 1804: 417.
    • Combination in Myrmica: Lepeletier de Saint-Fargeau, 1835: 185.
    • Combination in Micromyrma: Roger, 1862b: 258.
    • Combination in Tapinoma: Mayr, 1862: 651; Dalla Torre, 1893: 165.
    • Combination in Tapinoma (Micromyrma): Santschi, 1928e: 475.
    • Status as species: Latreille, 1802c: 269; Fabricius, 1804: 417; Lepeletier de Saint-Fargeau, 1835: 185; Smith, F. 1858b: 46; Roger, 1862b: 258; Roger, 1863b: 14; Mayr, 1863: 455; Mayr, 1876: 83; Forel, 1881: 3; Mayr, 1884: 31; Mayr, 1886c: 359; Emery, 1887a: 249; Forel, 1891b: 101 (redescription); André, 1892b: 51; Forel, 1893g: 352; André, 1893b: 152; Dalla Torre, 1893: 164; Emery, 1893d: 89; Emery, 1893e: 194; Emery, 1893f: 249; Emery, 1893g: 267; Forel, 1895a: 49; Forel, 1895b: 109; Forel, 1895e: 472; Emery, 1895f: 337; Pergande, 1896: 870; Mayr, 1897: 432; Emery, 1897d: 573; Forel, 1899a: 120; Forel, 1899c: 101; Emery, 1900d: 696; Emery, 1901f: 121; Forel, 1902h: 476; Rothney, 1903: 98; Bingham, 1903: 304; Wheeler, W.M. 1905b: 131; Emery, 1906c: 177; Wheeler, W.M. 1907a: 275; Forel, 1907e: 9; Wheeler, W.M. 1908a: 150; Forel, 1908e: 64; Wheeler, W.M. 1909c: 272; Wheeler, W.M. 1909d: 336, 342; Yano, 1910: 421; Wheeler, W.M. 1911a: 29; Wheeler, W.M. 1912a: 46; Forel, 1912k: 164; Emery, 1913a: 41; Forel, 1913f: 197; Forel, 1913k: 93; Wheeler, W.M. 1913b: 498; Wheeler, W.M. 1913d: 242; Wheeler, W.M. 1913e: 237; Santschi, 1914d: 377; Emery, 1914f: 420; Donisthorpe, 1915d: 342; Viehmeyer, 1916a: 142; Mann, 1916: 473; Wheeler, W.M. 1916d: 330; Crawley, 1916b: 375; Donisthorpe, 1918b: 167; Wheeler, W.M. 1919c: 275; Wheeler, W.M. 1919e: 100; Santschi, 1920h: 172; Mann, 1921: 473; Wheeler, W.M. 1922a: 924, 1034; Wheeler, W.M. 1922c: 15; Borgmeier, 1923: 101; Wheeler, W.M. 1923b: 4; Wheeler, W.M. 1924a: 110; Crawley, 1924: 403; Santschi, 1924c: 112; Mann, 1925b: 6; Mukerjee & Ribeiro, 1925: 207; Karavaiev, 1926d: 441; Stärcke, 1926: 118 (in key); Donisthorpe, 1927b: 395; Wheeler, W.M. 1927d: 7; Wheeler, W.M. 1927h: 99; Wheeler, W.M. 1928c: 30; Cheesman & Crawley, 1928: 522; Santschi, 1928c: 70; Wheeler, W.M. 1929f: 8; Wheeler, W.M. 1929g: 57; Karavaiev, 1930a: 213; Menozzi, 1930b: 114; Menozzi, 1930d: 328; Menozzi & Russo, 1930: 165; Smith, M.R. 1930a: 5; Wheeler, W.M. 1930h: 73; Donisthorpe, 1932c: 449; Wheeler, W.M. 1932a: 13; Wheeler, W.M. 1932d: 17; Wheeler, W.M. 1934a: 178; Wheeler, W.M. 1934h: 16; Donisthorpe, 1935: 634; Menozzi, 1935b: 200; Wheeler, W.M. 1935g: 36; Wheeler, W.M. 1936b: 204; Wheeler, W.M. 1937a: 23; Smith, M.R. 1937: 861; Teranishi, 1940: 59; Yasumatsu, 1940a: 68; Donisthorpe, 1941h: 60; Santschi, 1941: 277; Wheeler, W.M. 1942: 252; Menozzi, 1942: 178; Eidmann, 1944: 459, 469; Donisthorpe, 1948g: 139; Weber, 1948b: 85; Creighton, 1950a: 352; Smith, M.R. 1951a: 838; Azuma, 1951: 88; Chapman & Capco, 1951: 192; Smith, M.R. 1954c: 10; Kempf, 1961b: 520; Wilson, 1962c: 18; Linsley & Usinger, 1966: 175; Baltazar, 1966: 263; Wilson & Taylor, 1967: 80; Taylor, 1967b: 1094; Kempf, 1972a: 247; Alayo, 1974: 24 (in key); Smith, D.R. 1979: 1421; Collingwood, 1979: 36; Collingwood, 1985: 243; Deyrup, et al. 1989: 99; Wu, J. & Wang, 1992: 1311; Shattuck, 1994: 147; Douwes, 1995: 91; Bolton, 1995b: 400; Wu, J. & Wang, 1995: 116; Dorow, 1996a: 82; Tiwari, 1999: 83; Deyrup, et al. 2000: 300; Mathew & Tiwari, 2000: 342; Zhou, 2001b: 150; Blard, et al. 2003: 133; Deyrup, 2003: 47; Imai, et al. 2003: 84; Collingwood, et al. 2004: 486; Ghosh, et al. 2005: 10; Wild, 2007b: 25; Clouse, 2007b: 206; Don, 2007: 191; Framenau & Thomas, 2008: 59; Terayama, 2009: 199 (in key); Heterick, 2009: 58; Boer, 2010: 16; Collingwood, et al. 2011: 415; Csösz, et al. 2011: 58; Ellison, et al. 2012: 105; Sarnat & Economo, 2012: 37; Sharaf, Aldawood & El Hawagry, 2012a: 40 (in key); Borowiec, L. 2014: 168 (see note in bibliography); Wetterer, et al. 2016: 20; Radchenko, 2016: 92; Sharaf, Fisher, et al. 2017: 5; Deyrup, 2017: 176; Sharaf, Fisher, et al. 2018: 3; Guerrero, 2018: 499 (redescription).
    • Senior synonym of australis: Wilson & Taylor, 1967: 80; Shattuck, 1994: 148; Bolton, 1995b: 399; Radchenko, 2016: 92; Guerrero, 2018: 500.
    • Senior synonym of coronatum: Guerrero, 2018: 500.
    • Senior synonym of familiaris: Dalla Torre, 1893: 165; Forel, 1895b: 109; Forel, 1895e: 472; Pergande, 1896: 870; Forel, 1899c: 101; Wheeler, W.M. 1908a: 150; Emery, 1913a: 41; Wheeler, W.M. 1919c: 276; Wheeler, W.M. 1919e: 100; Wheeler, W.M. 1922a: 925; Borgmeier, 1923: 101; Creighton, 1950a: 352; Smith, M.R. 1954c: 10; Guerrero, 2018: 499.
    • Senior synonym of luffae: Guerrero, 2018: 500.
    • Senior synonym of malesiana: Guerrero, 2018: 500.
    • Senior synonym of nana: Emery, 1892b: 166; Dalla Torre, 1893: 165; Forel, 1895b: 109; Forel, 1895e: 472; Pergande, 1896: 870; Forel, 1899c: 101; Wheeler, W.M. 1908a: 150; Emery, 1913a: 41; Wheeler, W.M. 1919c: 275; Wheeler, W.M. 1919e: 100; Wheeler, W.M. 1922a: 925; Borgmeier, 1923: 101; Creighton, 1950a: 352; Smith, M.R. 1954c: 10; Radchenko, 2016: 92; Guerrero, 2018: 499.
    • Senior synonym of pellucida: Mayr, 1886c: 359; Forel, 1891b: 102; Dalla Torre, 1893: 165; Forel, 1895b: 109; Forel, 1895e: 472; Pergande, 1896: 870; Forel, 1899c: 101; Wheeler, W.M. 1908a: 150; Emery, 1913a: 41; Wheeler, W.M. 1919c: 275; Wheeler, W.M. 1919e: 100; Wheeler, W.M. 1922a: 925; Borgmeier, 1923: 101; Creighton, 1950a: 352; Smith, M.R. 1954c: 10; Radchenko, 2016: 92; Guerrero, 2018: 499.
    • Senior synonym of wallacei: unavailable name (proposed in synonymy), and also unnecessary (second) replacement name: Guerrero, 2018: 499.
  • australis. Tapinoma (Micromyrma) melanocephalum var. australis Santschi, 1928a: 53 (w.) VANUATU (date of publication 25.ii.1928), SAMOA (Upolu I., Savai’i I.), AMERICAN SAMOA (Tutuila I.).
    • [Also described as new, Tapinoma (Micromyrma) melanocephalum var. australe Santschi, 1928e: 475 (date of publication 30.viii.1928).]
    • Subspecies of melanocephalum: Wheeler, W.M. 1932c: 161; Wheeler, W.M. 1932d: 18; Wheeler, W.M. 1933f: 143; Wheeler, W.M. 1935g: 36; Wheeler, W.M. 1936f: 14.
    • Junior synonym of melanocephalum: Wilson & Taylor, 1967: 80; Shattuck, 1994: 148; Bolton, 1995b: 399.
  • coronatum. Tapinoma melanocephalum var. coronatum Forel, 1908b: 62 (w.) COSTA RICA.
    • Subspecies of melanocephalum: Emery, 1913a: 41; Kempf, 1972a: 247; Shattuck, 1994: 148; Bolton, 1995b: 400.
    • Junior synonym of melanocephalum: Guerrero, 2018: 500.
  • familiaris. Formica familiaris Smith, F. 1860b: 96 (w.) INDONESIA (Bacan I.).
    • [Junior primary homonym of Formica familiaris Smith, F. 1860a: 68.]
    • Status as species: Mayr, 1863: 414.
    • Replacement name: Formica domestica Smith, F. 1871a: 308.
    • Senior synonym of Formica wallacei Guerrero, 2018: 499. [Unavailable name (proposed in synonymy).] Unnecessary (second) replacement name for Formica familiaris Smith, F. 1860b: 96; junior synonym of Formica melanocephala Fabricius, 1793: 353: Guerrero, 2018: 499.
    • Junior synonym of melanocephalum: Forel, 1895e: 472; Pergande, 1896: 870; Forel, 1899c: 101; Guerrero, 2018: 499.
  • luffae. Neoclystopsenella luffae Kurian, 1955: 133, figs. 249-255 (m.) INDIA (National Capital Territory).
    • Combination in Tapinoma: Brown, 1988a: 337.
    • Status as species: Shattuck, 1994: 146; Bolton, 1995b: 400.
    • Junior synonym of melanocephalum: Guerrero, 2018: 500.
  • malesiana. Tapinoma melanocephalum var. malesiana Forel, 1913k: 93 (w.q.) INDONESIA (Sumatra, Java).
    • Subspecies of melanocephalum: Chapman & Capco, 1951: 192; Shattuck, 1994: 148; Bolton, 1995b: 400.
    • Junior synonym of melanocephalum: Guerrero, 2018: 500.
  • nana. Formica nana Jerdon, 1851: 125 (w.) INDIA (Karnataka; “found in all parts of India”).
    • [Duplicated in Jerdon, 1854b: 108.]
    • [Unresolved junior primary homonym of Formica nana Latreille, 1802c: 263 (Emery, 1921a: 26; Bolton, 1995b: 401).]
    • Status as species: Smith, F. 1858b: 15; Mayr, 1863: 418; Smith, F. 1871a: 303.
    • Junior synonym of melanocephalum: Emery, 1892b: 166; Forel, 1895e: 472.
  • pellucida. Myrmica pellucida Smith, F. 1857a: 71 (w.) SINGAPORE.
    • Status as species: Smith, F. 1858b: 124; Mayr, 1863: 434; Smith, F. 1871a: 324.
    • Junior synonym of melanocephalum: Mayr, 1886c: 359; Forel, 1895e: 472.

Type Material

  • Formica melanocephalum: Neotype (designated by Guerrero, 2018: 499), worker, Magdalena, Santa Marta, Calabazo, roadside to Pueblito-PNN Tayrona, Colombia, 11°16′57″N 74°01′26″W / 11.2825°N 74.0239°W / 11.2825; -74.0239, 22 January 2017, R. Guerrero, nest series, RJG01286 (ICNC-093716), Museo de Historia Natural; nest in dead branch on the roadside, secondary dry forest.
  • Tapinoma australis: Syntype, 23 workers, Santo Island, Hog Harbour, Vanuatu, The Natural History Museum, Naturhistorisches Museum, Basel.
  • Tapinoma melanocephalum coronatum: Lectotype (designated by Guerrero, 2018: 500), worker, Puntarenas, Costa Rica, CASENT0909775, Musee d'Histoire Naturelle Genève; middle specimen on the pin.
  • Tapinoma melanocephalum coronatum: Paralectotype (designated by Guerrero, 2018: 500), 2 workers, Puntarenas, Costa Rica, Musee d'Histoire Naturelle Genève; upper and lower specimen on the same pin as the lectotype.
  • Tapinoma melanocephalum malesiana: Lectotype (designated by Guerrero, 2018: 500), worker, Sumatra, Soengei Bamban, Indonesia, CASENT0909776, Musee d'Histoire Naturelle Genève.
  • Tapinoma melanocephalum malesiana: Paralectotype (designated by Guerrero, 2018: 500), 45 workers, 6 queens, Sumatra, Soengei Bamban, Indonesia, CASENT0909776, Musee d'Histoire Naturelle Genève.
  • Tapinoma melanocephalum malesiana: Paralectotype (designated by Guerrero, 2018: 500), 1 worker, Sumatra, Soengei Bamban, Indonesia, CASENT0909776, Naturhistorisches Museum, Basel.
  • Neoclystopsenella luffae: Holotype, male, Delhi, India, 2 July 1938, Imperial Entomologist, Forest Research Institute, Dehradun, India; On Luffa; additional types: Paratypes: 2 males, same data of holotype; 1 male same data of the holotype, except on cowpea and 5-VII-1938 (see Singh et al., 2021).

The following notes on F. Smith type specimens have been provided by Barry Bolton (details):

Formica familiaris

Holotype queen in Oxford University Museum of Natural History. Labelled “Mak.” (= Makassar, Sulawesi).

Myrmica pellucida

Four worker syntypes in Oxford University Museum of Natural History. Without a locality label on the specimens, but with a Smith det. label. Locality given in original description is Singapore.


Unless otherwise noted the text for the remainder of this section is reported from Guerrero, 2018.


Worker diagnosis. Bicolored small ants, with head and mesosoma black to brown, mandibles, antennae, legs and gaster light brown to pale yellow; third and fourth segment of the maxillary palps broad and flattened, spatulalike; basal end of the fifth segment inserted into the ventral face of the fourth segment.

Worker description (Figs 13–14): Head. Ovoid, longer than wide with slightly convex lateral margins; posterior margin variable, convex to straight, or sometimes slightly emarginated medially, in all cases, with rounded posterolateral corners. Mandibles subtriangular, straight outer margin over 2/3 length, and distally curved inwards; masticatory margin consisting of strong apical tooth, acute and longer than wide; slender subapical tooth, half length of the apical tooth; third triangular tooth half as long as subapical; fourth tooth slightly longer than third, followed by row of tight denticles decreasing in size approaching curvature leading to undifferentiated basal margin, the latter completely smooth or slightly crenulated. Palpal formula 6:4; maxillary palps with the first segment small and cylindrical, second segment conical and wider distally, sixth segment elongated, digitiform, rounded at tip; segments of labial palps relatively similar in size, 1st and 2nd segments wedge-shaped, 3rd subrectangular, last segment oval, longer than wide, with rounded tip. Clypeus broader than long; anteromedial margin with slight depression, and narrow, barely noticeable; margins lateral to depression relatively straight, those more lateral obliquely straight; posterior margin of clypeus convex, almost reaching base of frontal carinae. Frontal carinae diverging distally. Antennal scapes long, surpassing posterior margin of head by distance similar to length of pedicel, outer margins straight, slightly curved at both base and apex, wider near apical end; pedicel slightly conical with sub-parallel sides, third segment twice as wide as long, fourth-eleventh segments rectangular, much longer than wide, gradually elongating towards apex, last segment twice as long as others, rounded at tip. Eyes relatively large, oval, longer than wide, external margin relatively straight while the internal margin strongly convex, distance from the mandibular insertion approximately 1/2 maximum length of the eye.

Mesosoma. In lateral view, promesonotum strongly convex, forming nearly continuous line; metanotal sulcus broad and relatively deep; dorsopropodeum strongly convex, dome-shaped, at same level or slightly below level of promesonotum, declivitous face straight, inclined between 50–60º. Propodeal spiracle nearly circular, slightly longer than wide, opening laterally, well below half-length of lateropropodeum.

Metasoma. In lateral view, petiole short, flattened dorsal face with sharp apex inclined anteriorly; ventral lobe projecting, with straight anterior face and slightly curved posterior face. First tergum of gaster excavated anteriorly, as in other Tapinoma species. Body surface and pilosity. Cephalic surface opaque, finely covered with punctures and reticulations forming very small alveoli; abundant pubescence very short. Mandibles relatively smooth and slightly opaque, dorsal surface tenuously foveolate, covered with abundant short suberect hairs, and longer suberect hairs towards middle and basal portion; ventral face with long erect hairs. Dorsal face of clypeus covered with pubescence similar to that of cephalic capsule; anterior clypeal margin with two longer and thicker hairs, one on either side of medial depression, two golden relatively shorter setae than previously described hairs, above the clypeal margin are a pair of thick hairs directed anteriorly. Mesosoma completely covered with fine reticulations forming alveoli, relatively larger than those of head; surface rough and opaque, with dense whitish appressed hairs, very short and separated, sometimes imperceptible, giving appearance of glabrous surface, only visible in oblique-dorsal position. Legs with relatively fine granular and reticulated sculpture, relatively opaque, completely covered with dense whitish pubescence.

Petiole smooth and shiny, with few short, decumbent hairs near posteroventral margin. Gaster smooth to finely reticulated, relatively shiny, covered with fine dense golden yellowish pubescence; 5–6 erect hairs near posterior margin of fourth tergum, other terga without erect hairs; sternites with two erect hairs per segment. Coloration. Bicolored, variable in color tone. Head and mesosoma intense dark brown to brown with some regions yellowish brown on the propleura and the dorsum of the promesonotum. Mandibles, maxillary and labial palps, scapes, antennal segments, and legs pale yellow; masticatory margin reddish brown. Gaster brown to pale yellow as legs, but sometimes with contrasting brown bands on posterior margins of each tergum.

Neotype measurement: HL: 0.51 mm, HW: 0.45 mm, SL: 0.47 mm; WL: 0.44 mm; CI: 88; SI: 92.

Worker measurements (n= 316): HL: 0.46 (0.38–0.60) mm, HW: 0.41 (0.35–0.60) mm, SL: 0.43 (0.35–0.52) mm; WL: 0.39 (0.28–0.60) mm; CI: 89 (80–104); SI: 94 (82–105).


Gyne measurements (n= 20): HL: 0.52 (0.43–0.60) mm; HW: 0.51 (0.41–0.60) mm; SL: 0.45 (0.35–0.52) mm; WL: 0.74 (0.59–0.83) mm; CI: 93–104; SI: 81–93.

Gyne diagnosis: Cephalic capsule diverging posteriorly, much wider posterior to the compound eyes than anterior to them. Second segment of maxillary palps triangular, flattened, longer than wide, 3rd and 5th segments much broader and flattened, 3rd and 4th segment almost fused at the ends where they meet, the basal end of the 5th segment inserted into the ventral face of the 4th segment.

Gyne description (Figs 15–16): Head. Capsule cephalic square to rectangular (CI: 93–104), cuneiform; sides of the capsule relatively straight, lateral margin interrupted by the outer margin of the eyes; well-defined posterolateral corners, rounded, continuing to straight posterior cephalic margin, with shallow and wide concavity. Mandibles as in workers. Palpal formula 6:4; first segment of maxillary palps cylindrical and thickened, last segment thickened, digitiform, rounded at tip; segments of labial palps as in workers, but relatively broader and flattened. Dorsal face of clypeus as in workers, except for anterior margin relatively straighter and entire, with a slight mesal concavity. Frontal carinae divergent posteriorly. Scapes long, surpassing posterolateral corners of head by distance similar to length of pedicel; outer and inner margins straight in most of the length but slightly curved distally, wider mid-length; antennal segments as in workers. Compound eyes large, located at middle of length of cephalic capsule, relatively close to clypeal insertion; eyes located laterally, external margin breaking border of the cephalic capsule. Ocelli small, near level of posterior border of head; posterior ocelli near vertex but never breaking vertexal line.

Mesosoma. In profile, pronotum broad, propleura relatively broader than anterior part of pronotum; dorsal surface of mesonotum flattened; metanotum consisting of very thin strip, separated by deep mesonotal groove; dorsopropodeum without distinctive face, bending abruptly on a relatively inclined slope; mesopleuron divided into anepisternum and katepisternum, those divided by marked and relatively deep anapleural sulcus, anepisternum inverted U-shape, katepisternum subquadrate, almost as long as wide; propodeal spiracle circular, with an outer rim like a ring, directed posterad, and located on lateropropodeal face; metapleuron protruding, forming slight depression below propodeal spiracle. In dorsal view, mesoscutum strongly convex anteriorly, with parallel sides, diverging slightly posteriorly; mesonotal groove straight, slightly marked; mesoscutellum wider than long, posterior margin convex; dorsopropodeum restricted to very narrow band, posterior margin rounded bended abruptly on a slope, propodeum with the lateral margins, almost parallel.

Metasoma. Petiole relatively large, longer than wide, petiolar scale very low, restricted to small dorsal projection, inclined, directed anteriorly, with poorly developed lateral carinae; developed ventral lobe, most prominent posteriorly. Gaster as long as the mesosoma, flattened dorsally and slightly convex on ventral face. Body surface and pilosity. Surface of the head dense and finely punctate; with dense whitish decumbent hairs relatively separate, distance between hairs less than individual length. Dorsal face of mandibles densely covered with short golden hairs, on almost entire surface, except near area behind masticatory margin where they are long and inwardly directed. Maxillary palps covered with very short pilosity; labial palps with short erect and suberect hairs. Clypeus with pubescence as remainder of head, pattern of hairs on anterior margin as described in workers. Mesosoma opaque, abundantly and finely punctate; lateral regions of mesosoma covered with whitish pubescence, relatively longer than those of head, more abundant on katepisternum and the metapleuron; dorsum of the mesosoma covered with short pubescence, widely spaced, leaving surface partially exposed. Legs covered with short and decumbent hairs. Metasomal surface finely punctate as mesosoma, but relatively brighter; covered with abundant pubescence; fourth tergite with six long erect hairs near posterior margin, one central pair, and other two towards sides of the tergite.

Color. Head brown to dark brown, mandibles, maxillary and labial palps and antennae pale yellow; mesosoma bicolored, part of mesoscutum, mesoscutellum and legs yellowish brown to pale yellow, dorsum of pronotum and central portion of mesoscutum as spot, mesopleuron, inferior part of propodeum, and metapleuron brown. Petiole yellowish; gaster brown with yellowish brown bands on posterior margin


Male measurements (n= 17): HL: 0.44 (0.43–0.46) mm; HW: 0.42 (0.41–0.45) mm; SL: 0.39 (0.35–0.43) mm; WL: 0.66 (0.59–0.73) mm; CI: 93–104; SI: 81–93.

Male diagnosis: Third and fourth segment of the maxillary palps widened, oblong in shape, much longer than wide; the basal end of the 5th segment inserted into the ventral face of the 4th segment, like in the female caste.

Male description (Figs 17–18): Head. In dorsal view, head as long as broad, largest width posterior to compound eyes, with lateral margin convex, cephalic capsule converging anteriorly, posterolateral corners rounded, continuous with straight posterior cephalic margin; ventral surface of cephalic capsule relatively flattened. Mandibles long, semifalcate; external margin slightly sinuous, curving inwards distally; masticatory margin straight, with apical tooth relatively large, obliquely directed downwards, subapical denticle overhanging from margin, remainder of margin with very tight denticles, same size, continuing towards the undifferentiated basal margin. Palpal formula 6:4; 1st segment cylindrical, 2nd segment two times longer than first, cuneiform, 4th segment much wider than 3rd, 5th and 6th segments finger-shaped, the apical with rounded tip. Clypeus broad, relatively high, dorsal face outwardly projected; anteromedial border slightly concave, narrow, with rounded corners continuing toward relatively straight anterior margin, anterolateral margins inclined posteriorly; posterior margin convex but posterior vertex relatively flattened. Frontal carinae straight and semi-parallel at base, curved and diverging distally. Scape long, surpassing posterior cephalic margin by distance slightly less than distance between frontal carinae, scape with lateral margins parallel, outwardly curved from base to medial length, straight for rest of length, pedicel cuneiform, narrower at base, remainder of antennal segments cylindrical, longer than broad, last segment almost twice length of others. Compound eyes very large, oval, two times longer than broad, occupying almost entire lateral portion of head, breaking outline of lateral margins; anterior margin of eye separated from clypeal insertion by distance shorter than minimal width of scape. Ocelli large, located on wide base and relatively higher than dorsal surface of head, slightly below posterior cephalic vertex; anteromedial ocellus relatively smaller than lateral ocelli.

Mesosoma. In profile, propleura broad, flattened, with dorsal border thin, projecting dorsally; mesonotum anteriorly curved, with hump in lateral view, dorsal face relatively straight and high; mesopleuron divided into anepisternum and katepisternum, both delimited by wide and relatively deep anapleural sulcus; anepisternum inverted U-shaped, relatively wider than long, with dorsal margin slightly inclined posteriorly, anterior margin longer than posterior; katepisternum semiquadrate. Dorsopropodeum below level of mesonotum, with long surface, posteropropodeum slightly bulging decline of 90° toward base; propodeal spiracle oval, longer than broad, located on lateropropodeum, opening posteriorly. In dorsal view, anterior line of pronotum visible; mesoscutum slightly triangular with inferior margin truncated, longer than broad; dorsal face of propodeum visible.

Metasoma. In lateral view, petiole with petiolar scale well developed, anterior face relatively high and straight, forming angle with oblique dorsal face that descends posteriorly to declivity; ventral lobe developed, slightly convex. Cupula weakly sclerotized, easily collapsible when specimen pinned; telomeres dorsoventrally broad, obliquely truncated at apex; digitus long and narrow to tip, latter ventrally curved, cuspis located well inside genital capsule, semi-rectangular in lateral/medial view. Body surface and pilosity. Head surface opaque, softly punctulate; abundant, whitish, appressed hairs. Mandibles smooth, shining, bearing scattered short and long hairs, subdecumbent from base to behind masticatory margin; some hairs long, surpassing external margin. Anterior margin of clypeus with same pattern of hairs as described in worker; more lateral portions with relatively larger curved hairs; dorsal face densely hairy. Scape surface opaque, slightly punctate, covered by very short pilosity, with some hairs relatively longer and suberect in lateral margins; antennal segments with surface granulate, densely covered by short decumbent hairs, with some short suberect hairs.

Mesosoma relatively shining, with some areas notably opaque, puncticulate; surface covered by relatively short dense golden scattered pubescence, more abundant on katepisternum and metapleuron. Gaster reticulate, with small alveoli; covered by dense golden pubescence, but relatively sparse, leaving uncovered surface; telomeres densely covered by relatively long pilosity, surpassing margin.

Color. Ants completely yellowish brown, except for mandibles, scape and antennal segment, and legs which are whitish, contrasting with the remainder of body.

Larvae: Dolichoderine larvae described by Wheeler and Wheeler (1951), and examined in detail by Jesus et al. (2010).


  • 2n = 10, karyotype = 8M+2A (India) (Imai et al., 1984).
  • 2n = 10 (Indonesia) (Imai et al., 1985).
  • n = 5, 2n = 10 (USA) (Crozier, 1970a; Crozier, 1970b).


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