Tapinoma melanocephalum

AntWiki - Where Ant Biologists Share Their Knowledge
Jump to: navigation, search
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


The ghost ant, Tapinoma melanocephalum, is an ubiquitous indoor and outdoor pest throughout much of the tropics and subtropics, and an increasingly common indoor pest in temperate regions.

At a Glance • Polygynous  

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)

Distribution based on Regional Taxon Lists

Afrotropical Region: Cape Verde, Comoros, Guinea, 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, Paraguay, Puerto Rico, Suriname, Trinidad and Tobago.
Oriental Region: Bangladesh, Cambodia, India (type locality), Laos, Nicobar Island, Sri Lanka, Thailand.
Palaearctic Region: Afghanistan, Belgium, Canary Islands, China, Czech Republic, Finland, Hungary, Iberian Peninsula, 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


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.

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.

Associations with other Organisms

Other Insects

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


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.



Additional worker images can be found here




The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.

  • 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, Kapoor Collection; on Luffa, type material apparently lost (Guerrero, 2018: 500).

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.



  • Baltazar, C. R. 1966. A catalogue of Philippine Hymenoptera (with a bibliography, 1758-1963). Pac. Insects Monogr. 8: 1-488 (page 263, listed)
  • Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 401, Senior synonym of australe: new synonymy)
  • Collingwood, C. A. 1979. The Formicidae (Hymenoptera) of Fennoscandia and Denmark. Fauna Entomol. Scand. 8:1-174.
  • Crozier, R. H. 1970a. Karyotypes of twenty-one ant species (Hymenoptera: Formicidae), with reviews of the known ant karyotypes. Can. J. Genet. Cytol. 12: 109-128 (page 119, karyotype described)
  • Deyrup, M., Davis, L. & Cover, S. 2000. Exotic ants in Florida. Transactions of the American Entomological Society 126, 293-325.
  • Emery, C. 1887a [1886]. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Indo-Malese e dell'Australia. [part]. Ann. Mus. Civ. Stor. Nat. 24[=(2)(4): 209-240 (page 249, male described)
  • Emery, C. 1892c [1891]. Note sinonimiche sulle formiche. Bull. Soc. Entomol. Ital. 23: 159-167 (page 166, Senior synonym of nana)
  • Espadaler, X. 2007. The ants of El Hierro (Canary Islands). Pages 113-127 in R. R. Snelling, B. L. Fisher, and P. S. Ward, editors. Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson - 50 years of contributions. Memoirs of the American Entomological Institute, Gainesville, FL. 80:690 pp.
  • Fabricius, J. C. 1793. Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tome 2. Hafniae [= Copenhagen]: C. G. Proft, 519 pp. (page 353, worker described)
  • Forel, A. 1891c. Les Formicides. [part]. In: Grandidier, A. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (28e fascicule). Paris: Hachette et Cie, v + 237 pp. (page 102, queen described)
  • Forel, A. 1899f. Formicidae. [part]. Biol. Cent.-Am. Hym. 3: 81-104 (page 101, Senior synonym of familaris)
  • Guerrero, R.J. 2018. Taxonomic identity of the ghost ant, Tapinoma melanocephalum (Fabricius, 1793) (Formicidae: Dolichoderinae). Zootaxa 4410 (3): 497–510 (DOI 10.11646/zootaxa.4410.3.4).
  • Klimes, P. and J. Okrouhlik. 2015. Invasive ant Tapinoma melanocephalum (Hymenoptera: Formicidae): A rare guest or increasingly common indoor pest in Europe? European Journal of Entomology. 112:705-712. doi:10.14411/eje.2015.089
  • MacGown, J.A. 2017. First record of the Ghost Ant, Tapinoma melanocephalum (Hymenoptera: Formicidae), from Alabama. Transactions American Entomological Society 143: 533-535.
  • Mayr, G. 1862. Myrmecologische Studien. Verh. K-K. Zool.-Bot. Ges. Wien 12: 649-776 (page 651, Combination in Tapinoma)
  • Mayr, G. 1886c. Notizen über die Formiciden-Sammlung des British Museum in London. Verh. K-K. Zool.-Bot. Ges. Wien 36: 353-368 (page 359, Senior synonym of pellucida)
  • Obregon, R., M. R. Shaw, J. Fernandez-Haeger, and D. Jordano. 2015. Parasitoid and ant interactions of some Iberian butterflies (Insecta: Lepidoptera). Shilap-Revista De Lepidopterologia. 43:439-454.
  • Roger, J. 1862b. Beiträge zur Kenntniss der Ameisenfauna der Mittelmeerländer. II. Berl. Entomol. Z. 6: 255-262 (page 258, Combination in Micromyrma)
  • Santschi, F. 1928e. Nouvelles fourmis d'Australie. Bull. Soc. Vaudoise Sci. Nat. 56: 465-483 (page 475, Combination in Tapinoma (Micromyrma))
  • Sharaf, M.R., Fisher, B.L., Collingwood, C.A., Aldawood, A.S. 2017. Ant fauna (Hymenoptera: Formicidae) of the Socotra Archipelago (Yemen): zoogeography, distribution and description of a new species. Journal of Natural History 51, 317–378 (DOI 10.1080/00222933.2016.1271157).
  • Shattuck, S. O. 1994. Taxonomic catalog of the ant subfamilies Aneuretinae and Dolichoderinae (Hymenoptera: Formicidae). Univ. Calif. Publ. Entomol. 112:i-xix, 1-241. (page 148, see also)
  • Shi, Q. X., L. H. Lu, Y. Y. Lei, Y. R. He, and J. Chen. 2017. Gland Origin and Electroantennogram Activity of Volatile Compounds in Ghost Ants, Tapinoma melanocephalum (Hymenoptera: Formicidae) and Behavioral Response to (Z)-9-Nonadecene. Environmental Entomology. 46:1374-1380. doi:10.1093/ee/nvx164
  • Smith, D. R. 1979. Superfamily Formicoidea. Pp. 1323-1467 in: Krombein, K. V., Hurd, P. D., Smith, D. R., Burks, B. D. (eds.) Catalog of Hymenoptera in America north of Mexico. Volume 2. Apocrita (Aculeata). Washington, D.C.: Smithsonian Institution Pr (page 1421, see also)
  • Sorvari, J. 2002. Tapinoma melanocephalum (Fabricius, 1793) (Hymenoptera: Formicidae), an imported ant species new to Finland, with observations and a taxonomic note. Entomol. Gaz. 53: 269-270 (page 269, see also)
  • Wetterer, J. K. 2009. Worldwide spread of the ghost ant, Tapinoma melanocephalum (Hymenoptera: Formicidae). Myrmecol. News. 12:23-33.
  • Wheeler, G. C.; Wheeler, J. 1951. The ant larvae of the subfamily Dolichoderinae. Proc. Entomol. Soc. Wash. 53: 169-210 (page 197, larva described)
  • Wilson, E. O.; Taylor, R. W. 1967b. The ants of Polynesia (Hymenoptera: Formicidae). Pac. Insects Monogr. 14: 1-109 (page 80, Senior synonym of australis)
  • Zhou, A. M., B. Q. Kuang, and Y. R. Gao. 2015. Does the host plant affect the benefits from mutualisms? The invasive mealybug and ghost ant association. Ecological Entomology. 40:782-786. doi:10.1111/een.12258
  • Zima, J., Lebrasseur, O., Borovanska, M., Janda, J. 2016. Identification of microsatellite markers for a worldwide distributed, highly invasive ant species Tapinoma melanocephalum (Hymenoptera: Formicidae). European Journal of Entomology 113: 409–414 (doi: 10.14411/eje.2016.053).