Tapinoma melanocephalum

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)

Identification
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)

Distribution
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, Krakatau Islands, Malaysia, New Guinea, Niue, Philippines, Samoa, Singapore, Solomon Islands, Tokelau, Tonga, Vanuatu , 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, Costa Rica , 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, Laos, Maldives, Nicobar Island, Pakistan, 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.

Biology
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).

Foraging/Diet
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.

Other Insects
This ant has been observed tending larvae of (Obregon et al. 2015).

Chemistry
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.

Worker
Additional worker images can be found here

Nomenclature

 *  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 familiaris

Holotype queen in. Labelled “Mak.” (= Makassar, Sulawesi).

Myrmica pellucida

Four worker syntypes in. Without a locality label on the specimens, but with a Smith det. label. Locality given in original description is Singapore.

References based on Global Ant Biodiversity Informatics

 * Abe T., S. Yamane, and K. Onoyama. Ants collected on the Krakatau Islands 100 years after the great eruptions. Biogeography 14: 65-75.
 * Alcantara M. J., S. Modi, T. C. Ling, J. Monkai, H. Xu, S. Huang, and A. Nakamura. 2019. Differences in geographic distribution of ant species (Hymenoptera: Formicidae) between forests and rubber plantations: a case study in Xishuangbanna, China, and a global meta-analysis. Myrmecological News 29: 135-145.
 * Amarasinghe H. E. 2010. Species composition and nesting habits of ants in a hill-country home garden in Sri Lanka. Asian Myrmecology 3: 9-20.
 * André E. 1892. Voyage de M. Chaper à Bornéo. Catalogue des fourmis et description des espèces nouvelles. Mém. Soc. Zool. Fr. 5: 46-55.
 * Anh L. N., K. Ogata, and S. Hosoichi. 2010. Ants of agricultural fields in Vietnam (Hymenoptera: Formicidae). Bull. Inst. Trop. Agr., Kyushu Univ. 33: 1-11.
 * Baroni Urbani C. 1977. Katalog der Typen von Formicidae (Hymenoptera) der Sammlung des Naturhistorischen Museums Basel (2. Teil). Mitt. Entomol. Ges. Basel (n.s.) 27: 61-102.
 * Bharti H., Y. P. Sharma, M. Bharti, and M. Pfeiffer. 2013. Ant species richness, endemicity and functional groups, along an elevational gradient in the Himalayas. Asian Myrmecology 5: 79-101.
 * Bharti H., Y. P. Sharma, and A. Kaur. 2009. Seasonal patterns of ants (Hymenoptera: Formicidae) in Punjab Shivalik. Halteres 1(1): 36-47.
 * Borowiec L., and S. Salata. 2019. Next step in the invasion: Trichomyrmex mayri (Forel, 1902) new to the Philippines (Hymenoptera: Formicidae). Annals of the Upper Silesian Museum in Bytom Entomology 28(3): 1-3.
 * Chapman, J. W., and Capco, S. R. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia. Monogr. Inst. Sci. Technol. Manila 1: 1-327
 * Cheesman L. E., and W. C. Crawley. 1928. A contribution towards the insect fauna of French Oceania. - Part III. Formicidae. Ann. Mag. Nat. Hist. 10(2): 514-525.
 * Chen P., Y. Su, S. S. Rao, Y. F. Long, and C. H. Du. 2012. Study on the ant diversity in different Illicium verum stands. Journal of West China Forestry Science 41(1): 60-68.
 * Chen Y. Q., Q. Li, Y. L. Chen, Z. X. Lu, X. Y. Zhou. 2011. Ant diversity and bio-indicators in land management of lac insect agroecosystem in Southwestern China. Biodivers. Conserv. 20: 3017-3038.
 * Chen Y., C. W. Luo, H. W. Li, Y. J. Liu, H. F. Zheng, and F. C. Yang. 2013. Investigation of ant species and distribution on Wuliang Mountain. Journal of Henan Agricultural Sciences 42(5): 118-122.
 * Chen Y., C.-W. Luo, H. W Li, Z. H. Xu, Y. J. Liu, and S. J. Zhao. 2011. The investigation of soil ant resources on the West slope of Mt Ailao. Hubei Agricultural Sciences 50(7): 1356-1359.
 * Cheng D., Z. Chen, and S. Zhou. 2015. An analysis on the ant fauna of Jinzhongshan Nature Reserve in Gunagxi, China. Journal of Guangxi Normal University: Natural Science Edition 33(3): 129.137.
 * Crawley W.C. 1924. Ants from Sumatra, with biological notes by Edward Jacobson. Annals and Magazine of Natural History (9)13: 380-409
 * D'Cunha P., and V. M. Grover Nair. 2014. Ant fauna on the mangroves of Dakshina Kannada and Udupi districts, Karnataka, India. Journal of Ent. Res. 38(1): 59-66.
 * Dad J. M., S. A. Akbar, H. Bharti, and A. A. Wachkoo. 2019. Community structure and ant species diversity across select sites ofWestern Ghats, India. Acta Ecologica Sinica 39: 219–228.
 * Dhote J. 2012. Ant (Hymenoptera: Formicidae) records from Shri Shivaji science college campus, Amravati, India. Applied Research and Development Institute Journal 3(12): 114-117.
 * Dias R. K. S. 2002. Current knowledge on ants of Sri Lanka. ANeT Newsletter 4: 17- 21.
 * Dias R. K. S. 2006. Current taxonomic status of ants (Hymenoptera: Formicidae) in Sri Lanka. The Fauna of Sri Lanka: 43-52. Bambaradeniya, C.N.B. (Editor), 2006. Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. The World Conservation Union, Colombo, Sri Lanka & Government of Sri Lanka. viii + 308pp.
 * Dias R. K. S., K. R. K. A. Kosgamage, and H. A. W. S. Peiris. 2012. The Taxonomy and Conservation Status of Ants (Order: Hymenoptera, Family: Formicidae) in Sri Lanka. In: The National Red List 2012 of Sri Lanka; Conservation Status of the Fauna and Flora. Weerakoon, D.K. & S. Wijesundara Eds., Ministry of Environment, Colombo, Sri Lanka. p11-19.
 * Dias R. K. S., and K. R. K. A. Kosgamage. Systematics and community composition of foraging workers ants (Family: Formicidae) collected from three habitats in a dry zone region of Sri Lanka. Proceedings of the Annual Research Symposium 2008. Faculty of Graduate Studies, University of Kelaniya.
 * Dias R. K. S., and K. R. K. Anuradha Kosgamage. 2012. Occurrence and species diversity of ground-dwelling worker ants (Family: Formicidae) in selected lands in the dry zone of Sri Lanka. J. Sci. Univ. Kelaniya 7: 55-72.
 * Dias R. K. S., and R. P. K. C. Rajapaksa. 2016. Geographic records of subfamilies, genera and species of ants (Hymenoptera: Formicidae) in the four climatic zones of Sri Lanka: a review. J. Sci. Univ. Kelaniya 11(2): 23-45.
 * Dias R. K. S., and W. S. Udayakantha. 2016. Discovery of the Sri Lankan Relict Ant, Aneuretus simoni Emery (Formicidae, Aneuretinae) and the nest density of the species in a selected region of Meethirigala Forest Reserve, Sri Lanka. Asian Myrmecology 8: 1-8. DOI: 10.20362/am.008005
 * Dias R. K. S.; Perera K. A. M. 2011. Worker ant community observed by repeated sampling and information on endemic Aneuretus simoni Emery in the Gilimale Forest Reserve in Sri Lanka. Asian Myrmecology 4: 69-78.
 * Dias, R.K.S. 2006. Overview of ant research in Sri Lanka: 2000-2004. ANeT Newsletter 8:7-10
 * Emery C. 1893. Voyage de M. E. Simon aux îles Philippines (mars et avril 1890). Formicides. Annales de la Société Entomologique de France 62: 259-270.
 * Emery C. 1893. Voyage de M. E. Simon à l'île de Ceylan (janvier-février 1892). Formicides. Annales de la Société Entomologique de France 62: 239-258.
 * Emery C. 1901. Ameisen gesammelt in Ceylon von Dr. W. Horn 1899. Deutsche Entomologische Zeitschrift 1901: 113-122.
 * Emery C. Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei. Annali del Museo Civico di Storia Naturale 40: 661-722.
 * Emery, C. "Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei." Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova) (2) 20, no. 40 (1900): 661-722.
 * Fayle T.M., Bakker, L., Cheah, C., Ching, T.M., Davey, A., Dem, F., Earl, A., Huaimei, Y., Hyland, S., Johansson, B., Ligtermoet, E., Lim, R., Lin, L.K., Luangyotha, P., Martins, B.H., Palmeirim, A.F., Paninhuan, S., Rojas, S.K., Sam, L., Sam, P.T.T., Susanto, D., Wahyudi, A., Walsh, J., Weigl, S., Craze, P.G., Jehle, R., Metcalfe, D. & Trevelyan, R. 2011. A positive relationship between ant biodiversity (Hymenoptera: Formicidae) and rate of scavenger-mediated nutrient redistribution along a disturbance gradient in a south-east Asian rain forest. Myrmecological News 14: 5-12.
 * Fellowes J. R. 1999. Exotic ants in Asia: is the mainland at risk? The case of Hong Kong. Aliens 9: 5-6.
 * Forel A. 1895. Les Formicides de l'Empire des Indes et de Ceylan. Part V. J. Bombay Nat. Hist. Soc. 9: 453-472.
 * Forel A. 1911. Ameisen aus Ceylon, gesammelt von Prof. K. Escherich (einige von Prof. E. Bugnion). Pp. 215-228 in: Escherich, K. Termitenleben auf Ceylon. Jena: Gustav Fischer, xxxii + 262 pp.
 * Forel A. 1913. H. Sauter's Formosa-Ausbeute: Formicidae II. Arch. Naturgesch. (A)79(6): 183-202
 * [[Media:Forel 1913l.pdf|Forel A. 1913k. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien ausgeführt im Auftrage der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen. II. Ameisen aus Sumatra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren 1911-1912. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 36:1-148.]]
 * General D. M., and G. D. Alpert. 2012. A synoptic review of the ant genera (Hymenoptera, Formicidae) of the Philippines. Zookeys 200: 1-111.
 * Ghosh S. N., S. Sheela, B. G. Kundu, S. Roychowdhury, and R. N. Tiwari. 2006. Insecta: Hymenoptera: Formicidae. Pp. 369-398 in: Alfred, J. R. B. (ed.) 2006. Fauna of Arunachal Pradesh. (Part -2). [State Fauna Series 13.]. New Delhi: Zoological Survey of India, iv + 518 pp.
 * Ghosh S. N., S. Sheela, and B. G. Kundu. 2005. Ants (Hymenoptera: Formicidae) of Rabindra Sarovar, Kolkata. Records of the Zoological Survey of India. Occasional Paper 234: 1-40.
 * Guerrero R. J. 2018. Taxonomic identity of the ghost ant, Tapinoma melanocephalum (Fabricius, 1793) (Formicidae: Dolichoderinae). Zootaxa 4410(3): 497-510.
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Herwina H., N. Nasir, Jumjunidang, and Yaherwandi. 2013. The composition of ant species on banana plants with Banana Bunchy-top Virus (BBTV) symptoms in West Sumatra, Indonesia. Asian Myrmecology 5: 151-161.
 * Herwina H., and K. Nakamura. 2007. Ant species diversity study using pitfall traps in a small yard in Bogor Botanic garden, West Java, Indonesia. Treubia 35: 99-116.
 * Hosoichi S., S. H. Park, S. Tagane, M. M. Rahman, and K. Ogata. 2018. Domatia of the ant-plant Hydnophytum formicarum (Rubiaceae) captured as nests by two widespread ant species, Tapinoma melanocephalum and Tetramorium nipponense (Hymenoptera: Formicidae). Entomological News 127(5): 407-412.
 * Hu C.-H. 2006. Indigenized conservation and biodiversity maintenance on Orchid Island. PhD Thesis, graduate school of the University of Minnesota. 150 pages.
 * Imai H. T., M. Kubota, W. L. Brown, Jr., M. Ihara, M. Tohari, and R. I. Pranata. 1985. Chromosome observations on tropical ants from Indonesia. Annu. Rep. Natl. Inst. Genet. Jpn. 35: 46-48.
 * Ito, F.; Yamane, S.; Eguchi, K.; Noerdjito, W. A.; Kahono, S.; Tsuji, K.; Ohkawara, K.; Yamauchi, K.; Nishida, T.; Nakamura, K. 2001. Ant species diversity in the Bogor Botanic Garden, West Java, Indonesia, with descriptions of two new species of the genus Leptanilla (Hymenoptera, Formicidae). Tropics 10:379-404.
 * Jaitrong W., B. Guenard, E. P. Economo, N. Buddhakala, and S. Yamane. 2016. A checklist of known ant species of Laos (Hymenoptera: Formicidae). Asian Myrmecology 8: 1-32. DOI: 10.20362/am.008019
 * Jaitrong W., and T. Ting-Nga. 2005. Ant fauna of Peninsular Botanical Garden (Khao Chong), Trang Province, Southern Thailand (Hymenoptera: Formicidae). The Thailand Natural History Museum Journal 1(2): 137-147.
 * Jaitrong W.; Nabhitabhata, J. 2005. A list of known ant species of Thailand. The Thailand Natural History Museum Journal 1(1): 9-54.
 * Karmaly K. A.; S. Sumesh, T. P. Rabeesh, and L. Kishore. 2010. A checklist of ants of Thirunelli in Wayanad, Kerala. J. of the Bombay Natural History Society 107(1): 64-67.
 * Kishimoto-Yamata K., F. Hyodo, M. Matsuoka, Y. Hashimoto, M. Kon, T. Ochi, S. Yamane, R. Ishii, and T. Itioka. 2012. Effects of remnant primary forests on ant and dung beetle species diversity in a secondary forest in Sarawak, Malaysia. Journal of Insect Conservation DOI 10.1007/s10841-012-9544-6
 * Kosgamage K. R. K. A., and R. K. S. Dias 2008. Systematics and community composition of Foraging worker ants ((Order: Hymenoptera, Family: Formicidae) collected from three habitats on a dry zone region of Sri Lanka.Proceedings of Postgraduate Symposium of Kelaniya University. 115pp.
 * Lee T. L., and Y. S. Wei. 2005. Study for the temporal and spatial variation of the ant assemblages as the biological indicator in national parks. Journal of Animal and Veterinary Advances 4(4): 491-496.
 * Leong C. M., S. F. Shiao, and B. Guenard. 2017. Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau, one of the most heavily urbanized regions of the world. Asian Myrmecology 9: e009014.
 * Li Q., B. D. Hoffmann, Z. X. Lu, and Y. Q. Chen. 2017. Ants show that the conservation potential of afforestation efforts in Chinese valley-type savanna is dependent upon the afforestation method. Journal of Insect Conservation DOI 10.1007/s10841-017-0005-0
 * Li Q., Y. Chen, S. Wang, Y. Zheng, Y. Zhu, and S. Wang. 2009. Diversity of ants in subtropical evergreen broadleaved forest in Pu'er City, Yunnan. Biodiversity Science 17(3): 233-239.
 * Li Q., Z. Lu, Z. Wei, M. Yanyan, and F. Ping. 2015. Communities of ground-dwelling ants in different plantation forest in arid-hot valleys of Jinsha river, Yunnan Province, China. Scientia Silvae Sinicae 51(8): 134-142.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * Liu X. 2012. Taxonomy, diversity and spatial distribution characters of the ant family Formicidae (Insecta: Hymenoptera) in southeastern Tibet. PhD Thesis 139 pages
 * Lopwichan S., and S. Khachonpisitsak. 2015. Ant diversity in Nong Tha Yu Arboretum, Si Racha District, Chon Buri Province. Proceedings The 7 th National Science Research Conference. 30-31 March 2015. Naresuan University.
 * Lu Z. X., Y. Q. Chen, Q. Li, S. M. Wang, C. J. Liu, and W. Zhang. 2012. Effect of lac insect honeydew on the diversity of ground-dwelling ants in lac plantation. Chin. J. Appl. Ecol. 23(4): 1117-1122.
 * Lu Z., K. Li, N. Zhang, and Y. Chen. 2017. Diversity and indicator species of leaf-litter ants in Eucalyptus grandis plantations and secondary natural forests. Forest Research 29(4): 576-580
 * Lu Z., Y. Chen, Q. Li, S. Wang, C. Liu, and W. Zhang. 2012. Effect of population of Kerria yunnanensis on diversity of ground dwelling ant. Acta Ecologica Sinica 32(19): 6195-6202.
 * Lu Z., Y. Chen, W. Zhang, S. Wang, and Q. Li. 2013. Effects of facultative mutualism between ants and lac insects on the diversity of ant communities. Biodiversity Science 21(3): 343-351.
 * Lu Z., and Y. Chen. 2016. Effects of habitat on ant functional groups: a case study of Luchun County, Yunnan Province, China. Chinese Journal of Eco-Agriculture 24(5): 801-810.
 * Mezger D., and M. Pfeiffer. 2011. Influence of the arrival of Anoplolepis gracilipes (Hymenoptera: Formicidae) on the composition of an ant community in a clearing in Gunung Mulu National Park, Sarawak, Malaysia. Asian Myrmecology 4: 89-98.
 * Mohanraj P., M. Ali, and K. Veerakumari. 2010. Formicidae of the Andaman and Nicobar Islands (Indian Ocean: Bay of Bengal). Journal of Insect Science 10: Article 172
 * Mohanraj, P., M. Ali and K. Veenakumari. 2010. Formicidae of the Andaman and Nicobar Islands (Indian Ocean: Bay Of Bengal). Journal of Insect Science 10:172.
 * Mukherji D., and S. Ribeiro. 1925. On a collection of ants (Formicidae) from the Andaman Islands. Records of the Indian Museum 27: 205-209.
 * Musthak Ali T. M. 1982. Ant fauna (Hymenoptera: Formicidae) of Bangalore with observations on their nesting and foraging habits. Thesis Abstracts. Haryana Agricultural University 8: 370-371.
 * Ngoc Anh L., K. Ogata, and S. Hosoishi. 2010. Ants of agricultural fields in Vietnam (Hymenoptera: Formicidae). Bull. Inst. Trop. Agr. Kyushu Univ. 33: 1-11.
 * Nie L., M. Ni, D. Ning, H. Ran, B. Hassan, and Y. Xu. 2019. Comparing mechanisms of competition among introduced and resident ants in China: from behavior to trophic position (Hymenoptera: Formicidae). Myrmecological News 29: 125-133.
 * Pajni H. R., and R. K. Suri. 1978. First report on the Formicid fauna (Hymenoptera) of Chandigarh. Res. Bull. (Science) Punjab University 29: 5-12.
 * Parui A. K., S. Chatterjee, and P. Basu. 2015. Habitat characteristics shaping ant species assemblages in a mixed deciduous forest in Eastern India. Journal of Tropical Ecology: 1-14. doi:10.1017/S0266467415000036
 * Pfeiffer M., D. Mezger, and J. Dyckmans. 2013. Trophic ecology of tropical leaf litter ants (Hymenoptera: Formicidae) - a stable isotope study in four types of Bornean rain forest. Myrmecological News 19: 31-41.
 * Pfeiffer M., and D. Mezger. 2012. Biodiversity Assessment in Incomplete Inventories: Leaf Litter Ant Communities in Several Types of Bornean Rain Forest. PLoS ONE 7(7): e40729. doi:10.1371/journal.pone.0040734
 * Pfeiffer M., and D. Mezger. 2012. Biodiversity Assessment in Incomplete Inventories: Leaf Litter Ant Communities in Several Types of Bornean Rain Forest. PLoS ONE 7(7): e40729. doi:10.1371/journal.pone.0040846
 * Pfeiffer M.; Mezger, D.; Hosoishi, S.; Bakhtiar, E. Y.; Kohout, R. J. 2011. The Formicidae of Borneo (Insecta: Hymenoptera): a preliminary species list. Asian Myrmecology 4:9-58
 * Pfeiffer, M., H. Cheng Tuck, and T. Chong Lay. 2008. Exploring arboreal ant community composition and co-ccurrence patterns in plantations of oil palm Elaeis guineensis in Borneo and Peninsular Malaysia. Ecography 31(1): 21-32.
 * Philpott S.M., P. Bichier, R.A. Rice, and R. Greenberg. 2008. Biodiversity conservation, yield, and alternative products in coffee agroecosystems in Sumatra, Indonesia. Biodivers. Conserv. 17: 1805-1820. Data obtained from Stacy Philpott
 * Raci N., C. Sravanthy, C. Sammaiah, and M. Thirupahaiah. 2015. Biodiversity of ants (Insecta-Hymenoptera) in agroecosystem and grass land in Jammikunta, Karimnagar District, Telangana, India. Journal ofEnvironment 4(1): 11-16.
 * Rajagopal T., S. P. Sevarkodiyone, and A. Manimozhi. 2005. Ant diversity in some selected localities of Sattur Taluk, Virudhunagar district, Tamil Nadu. Zoos' Print Journal 20(6): 1887-1888.
 * Rajan P. D., M. Zacharias, and T. M. Mustak Ali. 2006. Insecta: Hymenoptera: Formicidae. Fauna of Biligiri Rangaswamy Temple Wildlife Sanctuary (Karnataka). Conservation Area Series, Zool. Surv. India.i-iv,27: 153-188.
 * Ramesh T., K. J. Hussain, K. K. Satpathy, M. Selvanayagam, and M. V. R. Prasad. 2010. Diversity, distribution and species composition of ants fauna at Department of Atomic Energy (DAE) Campus Kalpakkam, South India. World Journal of Zoology 5 (1): 56-65.
 * Rizali A., A. Rahim, B. Sahari, L.B. Prasetyo, and D. Buchori. 2011. Impact of invasive ant species in shaping ant community structure on small islands in Indonesia. Jurnal Biologi Indonesia 7(2): 221-230.
 * Rizali A., M. M. Bos, D. Buchori, Sk. Yamane, and C. H. Schulze. 2008. Ants in tropical urban habitats: the myrmecofauna in a densely populated area of Bogor, West Java, Indonesia. Hayati Journal of Biosciences 15(2): 77-84.
 * Rizali A., M.M. Bos, D. Buchori, Sk. Yamane, C. Hans, and J. Schulze. 2008. Ants in tropical urban habitats: the myrmecofauna in a densely populated area of Bogor, West Java, Indonesia. Hayati Journal of Biosciences 77-84.
 * Rizali A., Y. Clough, D. Buchori, M. L . A. Hosang, M. M. Bos, and T. Tscharntke. 2012. Long-term change of ant community structure in cacao agroforestry landscapes in Indonesia. Insect Conservation and Diversity doi: 10.1111/j.1752-4598.2012.00219.x
 * Sakchoowong W., W. Jaitrong, and K. Ogata. 2009. Comparison of ground-ant diversity between natural forests and disturbed forests along a natural gas pipeline transect in Thong Pha Phum National Park, Kanchanaburi province. Kasetsart J. (Nat. Sci) 43: 64-73.
 * Savitha S., N. Barve, and P. Davidar. 2008. Response of ants to disturbance gradients in and around Bangalore, India. Tropical Ecology 49(2): 235-243.
 * Shattuck S. O. 1994. Taxonomic catalog of the ant subfamilies Aneuretinae and Dolichoderinae (Hymenoptera: Formicidae). University of California Publications in Entomology 112: i-xix, 1-241.
 * Shukla R. K., H. Singh, N. Rastogi, and V. M. Agarwal. 2013. Impact of abundant Pheidole ant species on soil nutrients in relation to the food biology of the species. Applied Soil Ecology 71: 15 23.
 * Sitthicharoenchai D., and N. Chantarasawat. 2006. Ant species diversity in the establishing area for Advanced Technology Institute at Lai-Nan Sub-district, Wiang Sa district, Nan Province, Thailand. The Natural History Journal of Chulalongkorn University 6(2): 67-74
 * Skarbek C. J., M. Noack, H. Bruelheide, W. Hardtle, G. von Oheimb, T. Scholten, S. Seitz, M. Staab. 2019. A tale of scale: plot but not neighbourhood tree diversity increases leaf litter ant diversity. Journal of Animal Ecology DOI: 10.1111/1365-2656.13115
 * Song Y., Z. H. Xu, C. L. Li, Y. Q. Hao, and H. B. Li. 2014. Ant communities of Nangun River Nature Reserve in Yunnan. Journal of West China Forestry Science 45(5): 93-100.
 * Song Y., Z. Xu, C. Li, N. Zhang, L. Zhang, H. Jiang, and F. Mo. 2013. An Analysis on the Ant Fauna of the Nangun river Nature Reserve in Yunnan, China. Forest Research 26(6): 773-780.
 * Sonune B. V., and R. J. Chavan. 2016. Distribution and diversity of ants (Hymenoptera: Formicidae) around Gautala Autramghat Sanctuary, Aurangabad Maharashtra, India. Journal of Entomology and Zoology Studies 4(2): 361-364.
 * Staab M., J. Methorst, J. Peters, N. Bluthgen, and A. M. Klein. 2017. Tree diversity and nectar composition affect arthropod visitors on extrafloral nectaries in a diversity experiment. Journal of Plant Ecology 10(1): 210-212.
 * Staab M., N. Bluthgen, and A. M. Klein. 2014. Tree diversity alters the structure of a tri-trophic network in a biodiversity experiment. Oikos doi: 10.1111/oik.01723
 * Tak N. 1995. Studies on ants (Formicidae) of Rajasthan - 1 Jodhpur. Hexapoda 7(1): 17-28.
 * Tak N. 2000. Studies on ants (Formicidae) of Rajasthan--II Dungarpur. Entomon 25: 47-54.
 * Tak N. 2008. Ants of Rajasthan. Conserving Biodiversity of Rajasthan Zool. Surv. India. 149-155.
 * Tak N. 2009. Ants Formicidae of Rajasthan. Records of the Zoological Survey of India, Occasional Paper No. 288, iv, 46 p
 * Tak N. 2010. Insecta: Hymenoptera: Formicidae. Zool. Surv. India, Fauna of Ranthambore National Park, Conservation Area Series 43: 133-144.
 * Tak N., N. S. Rathore, and S. Kumar. 2007. Insecta: Hymenoptera. Fauna of Pichhola lake (Rajasthan). Wetland Ecosystem series. Zool. Surv. India. 8 : 127-130.
 * Tak N., and N. S. Rathore. 1996. Ant (Formicidae) fauna of the Thar Desert. Pp. 271-276 in: Ghosh, A. K.; Baqri, Q. H.; Prakash, I. (eds.) 1996. Faunal diversity in the Thar Desert: gaps in research. Jodhpur: Scientific Publishers, xi + 410 pp.
 * Tak N., and N. S. Rathore. 2004. Insecta: Hymenoptera: Formicidae. State Fauna Series 8: Fauna of Gujarat. Zool. Surv. India. Pp. 161-183.
 * Tak N., and S. I. Kazmi. 2013. On some ants (Insecta: Hymenoptera: Formicidae) from Nagaland, India. Rec. zool. Surv. India: 113(1): 169-182.
 * Tak N., and S. L. Kazmi. 2011. On a collection of Insecta: Hymenoptera: Formicidae from Uttarakhand. Rec. zool. Surv. India : 111(2) : 39-49.
 * Tak, N. 2009. Ants (Hymenoptera: Formicidae) of the Thar Desert of Rajasthan and Gujarat. in C. Sivaperuman et al. (eds.), Faunal Ecology and Conservation of the Great Indian Desert
 * Terayama M. 2009. A synopsis of the family Formicidae of Taiwan (Insecta: Hymenoptera). Research Bulletin of Kanto Gakuen University. Liberal Arts 17:81-266.
 * Terayama, M. 2009. A synopsis of the family Formicidae of Taiwan (Insecta; Hymenoptera). The Research Bulletin of Kanto Gakuen University 17: 81-266.
 * Trainor C.R. and A.N. Andersen. 2010. The ant fauna of Timor and neighbouring islands: potential bridges between the disjunct faunas of South East Asia and Australia. Australian Journal of Zoology 58: 133-144.
 * Trainor C.R., and A.N. Andersen. 2010. The ant fauna of Timor and neighbouring islands: potential bridges between the disjunct faunas of South East Asia and Australia. Australian Journal of Zoology 58: 133-144.
 * Varghese T. 2004. Taxonomic studies on ant genera of the Indian Institute of Science campus with notes on their nesting habits. Pp. 485-502 in : Rajmohana, K.; Sudheer, K.; Girish Kumar, P.; Santhosh, S. (eds.) 2004. Perspectives on biosystematics and biodiversity. Prof. T.C. Narendran commemoration volume. Kerala: Systematic Entomology Research Scholars Association, xxii + 666 pp.
 * Wang C. and Wu J. 1992. Ants of the Jianfengling forest region in Hainan Province (Hymenoptera: Formicidae). Scientia Silvae Sinicae 28: 561-564.
 * Wang W. R., S. Q. Zhang, and Z. H. Xu. 2012. A faunistic and taxonomic study of ants (Hymenoptera: Formicidae) in Shenzhen Municipality. Journal of Southwest Forestry University 32(1): 64-73.
 * Wang W., S. Zhang, and Z Xu. 2012. Distribution patters of ant species in Shenzhen City. Journal of Southwest Forestry University 32(3): 70-74.
 * Wang W., S. Zhang, and Z. Xu. 2012. Distribution Patterns of Ant Species in Shenzhen City. Journal of Southwest Forestry University 32(3): 69-74.
 * Watanasit S., S. Sonthichai, and N. Noon-anant. 2003. Preliminary survey of ants at Tarutao National Park, Southern Thailand. Songklanakarin J. Sci. Technol. 25(1) : 115-122
 * Way M. J., and B. Bolton. 1997. Competition between ants for coconut palm nesting sites. Journal of Natural History 31: 439-455.
 * Wheeler W. M. 1909. Ants of Formosa and the Philippines. Bulletin of the American Museum of Natural History 26: 333-345.
 * Wheeler W. M. 1913. Zoological results of the Abor Expedition, 1911-1912, XVII. Hymenoptera, II: Ants (Formicidae). Records of the Indian Museum 8: 233-237.
 * Wheeler W. M. 1919. The ants of Borneo. Bulletin of the Museum of Comparative Zoology 63:43-147.
 * Wheeler W. M. 1923. Chinese ants collected by Professor S. F. Light and Professor A. P. Jacot. American Museum Novitates 69: 1-6.
 * Wheeler W. M. 1927. Ants collected by Professor F. Silvestri in Indochina. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 20: 83-106.
 * Wheeler W. M. 1928. Ants collected by Professor F. Silvestri in China. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 22: 3-38.
 * Wheeler W. M. 1929. Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 24: 27-64.
 * Wheeler W. M. 1929. Some ants from China and Manchuria. American Museum Novitates 361: 1-11.
 * Wheeler W. M. 1930. A list of the known Chinese ants. Peking Natural History Bulletin 5: 53-81.
 * Wheeler W. M. 1930. Formosan ants collected by Dr. R. Takahashi. Proceedings of the New England Zoological Club 11: 93-106.
 * Wheeler W. M. 1937. Additions to the ant-fauna of Krakatau and Verlaten Island. Treubia 16: 21-24.
 * Wu B., Y. Lu, G. Liang, and L. Zeng. 2010. Influence of the red inported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae) on the diversity of ant communities in a newly infested longan orchard and grass areas nearby. Acta Ecologica Sinica 30(8): 2075-2083.
 * Wu B., Y. Lu, L. Zeng, and G. Liang. 2008. Influences of Solenopsis invicta Buren invasion on the native ant communities in different habitats in Guangdong. Chin. J. Appl. Ecol. 19(1): 151-156.
 * Wu Z. W., X. D. Bi, X. He, Z. X. Lu, L. J. Wei, and Y. Q Chen. 2015. Impact of continuous fire disturbance on ground-dwelling ant communities in arid-hot valleys of Panzhihua, Sichuan. Journal of Yunnan University 37(3): 467-474.
 * Xu F. F., and J. Chen. 2009. Comparison of the Differences in Response to the Change of the Extrafloral Nectar-ant-herbivore Interaction System Between a Native and an Introduced Passiflora Species. Acta Botanica Yunnanica 31(6): 543-550.
 * Xu Z. H., B. L. Yang, and G. Hu. 1999. Formicidae ant communities in fragments of montane rain forest in Xishuangbanna, China. Zoological Research 20(4): 288-293.
 * Xu Z., Zeng G., Liu T.-Y. and He Y.-F. 1999. [A study on communities of Formicidae ants in different subtypes of vegetation in Xishuangbanna District of China.] Zoological Research 20: 118-125
 * Yamane S. 2013. A Review of the ant fauna of the Krakatau Islands, Indonesia. Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist. Ser: A, 11: 1-66
 * Yamane Sk. 2005. Krakatau in 1982, and the commencement of myrmecological research. The nature and Insects (Konchu to shizen) 40: 27-33.
 * Yasumatsu, K. 1940. Matériaux pour servir à la faune myrmécologique des îles de Yaeyama. Mushi 13: 67-70
 * Zhang C., Z. Xu, N. Yu, Q. He, and X. Liu. 2012. Distribution Patterns of Ant Species on East Slope of Mount Demola and Zayu Valley in Southeastern Tibet. Journal of Northeast Forestry University 40(3): 87-92
 * Zhang N. N., Y. Q. Chen, Z. X. Lu, W. Zhang, and K. L. Li. 2013. Species diversity, community structure difference and indicator species of leaf-litter ants in rubber plantations and secondary natural forests in Yunnan, southwestern China. Acta Entomologica Sinica 56(11): 1314-1323.
 * Zhang R. J., L. W. Liang, and S. Y. Zhou. 2014. An analysis on the ant fauna of Nonggang Nature Reserve in Guangxi, China. Journal of Guangxi Normal university: Natural Science Edition 32(3): 86-93.
 * Zhang W., G. Liu, P. Zhong, and S. Zhang. 2014. Investigation of Formicidae in Luofushan Mountain. Journal of Huizhou University 34(3): 46-50.
 * Zhang W., and S. Zhou. 2016. An investigation on Formicidae species of Nanling National Park. Journal of Huizhou University 36(3): 27-30.
 * Zhang Z., M. Cao, X. Yang, X. Deng, and Y. She. 2000. A study on species diversity of ant in fragments of seasonal rain forest of Xishuangbanna, China. Zoological Research 21(1): 70-75.
 * Zhou S.-Y. 2001. Ants of Guangxi. Guangxi Normal University Press, Guilin, China, Guilin, China. 255 pp.
 * Zryanin V. A. 2011. An eco-faunistic review of ants (Hymenoptera: Formicidae). In: Structure and functions of soil communities of a monsoon tropical forest (Cat Tien National Park, southern Vietnam) / A.V. Tiunov (Editor). – M.: KMK Scientific Press. 2011. 277 р.101-124.
 * Zryanin V. A., and M. V. Mokrousov. 2015. Contribution to the ant fauna of Lombok Island. Proceedings of the 10th ANeT International Conference, 23-26 October 2015, University of Kelaniya, Sri Lanka. 34