Dolichoderinae

The subfamily Dolichoderinae is one of four major species-rich subfamilies of ants, the others being Formicinae, Myrmicinae and Ponerinae. It is found world-wide and includes some of the world's most invasive species, including Linepithema humile (the Argentine ant), Tapinoma melanocephalum, Technomyrmex albipes and Technomyrmex difficilis.

Most species of Dolichoderinae are general predators or scavengers. Many also tend Hemiptera to collect honeydew or are associated with caterpillars. Nests are found in a wide variety of locations, including in the soil, under rocks and other objects, in rotten and living wood, in termite mounds, and in cracks between rocks. Some species (in the genus Arnoldius) are thought to establish their nests by new queens invading the nests of other ants, killing the queen, and using the captured workers to help raise her own offspring.

Species of Dolichoderinae can be found in most regions of the world and in all major habitats. They also have a rich fossil history, with 20 fossil genera known, the third richest after Myrmicinae (38 genera) and Formicinae (31 genera). The relationships among these genera have been well documented (see Phylogeny of Dolichoderinae).

The Dolichoderines lack a sting, but are armed with defensive compounds produced by the anal gland - a structure unique to the subfamily. This is the source of the pungent, and often unpleasant odors produced by many species when they are disturbed, crushed or otherwise annoyed. In many species colonies may be polygynous and/or polydomous. Some are predaceous, but most appear to be generalized scavengers with a strong liking for carbohydrates taken directly or indirectly from plant sources. In addition to founding colonies via single newly mated queens, some produce new colonies by budding or fission. Most Dolichoderines are free-living; a few are social parasites on other members of their subfamily.

Identification
The mesosoma is attached to the gaster by a single distinct segment, the petiole. The gaster is smooth, without constrictions between the segments. The sting is absent and the tip of the gaster is slit-like and without a circular opening (an acidopore).

Species of Dolichoderinae are most often confused with species of the subfamily Formicinae because both have a single segmented petiole, lack a sting and are often similar in overall body size and shape This is especially true for the smaller species such as those in Doleromyrma and Tapinoma. However, dolichoderines can always be separated from formicines because the tip of the gaster has a slit-like opening while all formicines have a small circular opening.

Males: Boudinot (2015) - The Dolichoderinae are uniquely identified by the telomere, which is strongly reduced and does not extend anteroventrad the basimere. Males of the subfamily are further identified by the following combination of characters: oblique mesopleural sulcus present; seven or fewer closed cells present on forewing; jugal lobe absent; petiolar peduncle short or absent; abdominal segment III unpetiolated; abdominal segment IV without cinctus between pre- and postsclerites; abdominal sternum IX unpronged and edentate. Additional characters for distinguishing males of the Dolichoderinae from the Aneuretinae and Formicinae are indicated in couplets 19 and 20 of the key to subfamilies key.

Shattuck (1992b): Seven characters were found which suggest the subfamily is monophyletic. These are

(1) Loss of pupal cocoons

(2) Reduction of the larval neck

(3) Reduced number of larval hairs

(4) Lightly sclerotized larval mandibles

(5) Larval maxillary palp and galea reduced to sensilla

(6) Small larval sericteries

(7) Cyclopentanoid monoterpene production.

Nomenclature

 * [subfamily of Formicidae]
 * Dolichoderidae Forel, 1878: 364. Type-genus: Dolichoderus Lund, 1831a: 130.

Taxonomic History

 * Dolichoderinae as family: Emery, 1894g: 378 [Dolichoderidae]; Ashmead, 1905b: 384 [Dolichoderidae]; Novák & Sadil, 1941: 94 [Dolichoderidae]; Bernard, 1951: 1071 [Dolichoderidae]; Bernard, 1953b: 255 [Dolichoderidae].
 * Dolichoderinae as tribe of Formicidae: André, 1882a: 127 [Dolichoderidae].
 * Dolichoderinae as subfamily of Formicidae: Forel, 1878: 364 [Dolichoderidae]; Emery & Forel, 1879a: 454 [Dolichoderidae]; Nasonov, 1889: 26 [Dolichoderidae]; Forel, 1892j: 220 [Dolichoderidae]; Forel, 1893a: 165; Dalla Torre, 1893: 156; Forel, 1895b: 107 [Dolichoderidae]; Emery, 1895j: 771 [subfamily spelled Dolichoderini]; Emery, 1896e: 186; Forel, 1899c: 98; Bingham, 1903: 288; Ruzsky, 1905b: 102; Wheeler, W.M. 1910g: 142; Emery, 1913a: 2; Arnold, 1915: 143; Wheeler, W.M. 1915h: 71; Wheeler, 1915g: 811 [Dolichoderides]; Arnold, 1915: 143; Donisthorpe, 1915d: 178; Forel, 1917: 247; Escherich, 1917: 2 [Dolichoderini]; Bondroit, 1918: 86 [Dolichoderitae]; Wheeler, W.M. 1920: 53; Wheeler, W.M. 1922a: 199; Borgmeier, 1923: 80; Karavaiev, 1936: 163; Smith, M.R. 1951a: 833; Clark, 1951: 16; Brown, 1954e: 29; Kempf, 1972a: 266; Wheeler, G.C. & Wheeler, J. 1972a: 41; Brown, 1973b: 169; subsequent authors; Shattuck, 1992c: 20.
 * Dolichoderinae as formicomorph subfamily of Formicidae: Bolton, 2003: 18, 80.
 * Dolichoderinae as formicoid subfamily of Formicidae: Brady, et al. 2006: 18173; Moreau, et al. 2006: 102.
 * Dolichoderinae as formicoid dolichoderomorph subfamily of Formicidae: Ward, 2007a: 556.

Taxonomic References
Forel, 1878: 364, 380 (diagnosis, genera); Dalla Torre, 1893: 156 (catalogue); Emery, 1895j: 771 (synoptic classification); Emery, 1896e: 186 (genera key); Handlirsch, 1907: 869 (*fossil taxa catalogue); Wheeler, W.M. 1910g: 142 (diagnosis); Emery, 1913a: 2, 6 (diagnosis, tribe key, catalogue); Emery, 1913a: 17 (Tapinomini diagnosis, genera key, catalogue); Arnold, 1915: 144, (diagnosis); Gallardo, 1916a: 3 (diagnosis); Forel, 1917: 247 (synoptic classification); Forel, 1921c: 136 (diagnosis); Wheeler, W.M. 1922a: 199, 687, 688 (diagnosis, tribe key, Tapinomini genera key); Brown & Nutting, 1950: 127 (venation, phylogeny); Brown, 1954e: 29 (phylogeny); Pavan, 1955: 135 (gastral organs); Eisner, 1957: 453 (proventriculus morphology); Bernard, 1967: 246 (diagnosis); Gotwald, 1969: 118 (mouthparts morphology); Wheeler, G.C. & Wheeler, J. 1972a: 41 (diagnosis); Brown, 1973b: 169 (genera and distribution); Wheeler, G.C. & Wheeler, J. 1976b: 61 (larvae, review and synthesis); Snelling, R.R. 1981: 401 (synoptic classification); Dazzini Valcurone & Fanfani, 1985: 1 (gastral glands); Wheeler, G.C. & Wheeler, J. 1985: 258 (synoptic classification); Billen, 1986: 173 (Dufour's gland); Billen, 1987: 278 (abdominal glands); Baroni Urbani & Wilson, 1987: 1 (*fossil Leptomyrmecini); Dlussky & Fedoseeva, 1988: 77 (synoptic classification); Hölldobler & Wilson, 1990: 9 (synoptic classification, genera keys); Baroni Urbani, et al. 1992: 316 (phylogeny); Shattuck, 1992b: 199 (higher classification, phylogeny); Shattuck, 1992c: 20 (revision of subfamily, genera key); Shattuck, 1994: 3 (catalogue); Bolton, 1994: 22 (diagnosis, synoptic classification, genera keys); Shattuck, 1995: 217 (phylogeny, genera); Bolton, 1995a: 1038 (census); Bolton, 1995b: 10 (catalogue); Wenseleers, Schoeters, et al., 1998: 121 (cloacal gland); Brandão, Baroni Urbani, et al. 1999: 411 (phylogeny, genera); Chiotis, et al. 2000: 108 (phylogeny, genera); Dlussky & Rasnitsyn, 2002: 416 (diagnosis for impression fossils); Bolton, 2003: 18, 80 (diagnosis, synopsis); Dubovikoff, 2005a: 91 (tribes of Dolichoderinae); Brady, et al. 2006: 18173 (phylogeny); Moreau, et al. 2006: 102 (phylogeny); Ward, et al. 2010: 342 (phylogeny, tribe-rank classification); Keller, 2011: 1 (morphology, phylogeny); Boudinot, 2015: 50 (diagnosis); Fisher & Bolton, 2016: 43 (diagnosis).

Regional and National Faunas with Keys
André, 1882a: 127 (Europe and Algeria); Nasonov, 1889: 50 (Russia); Forel, 1891b: 9 (Madagascar genera); Forel, 1895e: 460 (India and Sri Lanka); Bingham, 1903: 288 (India, Sri Lanka and Burma); Ruzsky, 1905b: 102 (Russian Empire); Bondroit, 1910: 488 (Belgium); Wheeler, W.M. 1910g: 560 (North America genera); Stitz, 1914: 77 (Central Europe); Gallardo, 1915: 34 (Argentina genera); Forel, 1915d: 40 (Switzerland); Arnold, 1915: 145 (South Africa); Donisthorpe, 1915d: 178 (Britain); Gallardo, 1916a: 12 (Argentina); Emery, 1916b: 208 (Italy); Wheeler, W.M. 1916m: 589 (U.S.A., Connecticut); Bondroit, 1918: 86 (France and Belgium); Gallardo, 1919b: 253 (Argentina genera); Soudek, 1922: 58 (Czechoslovakia); Stärcke, 1926: 117 (Netherlands); Karavaiev, 1927c: 271 (Ukraine); Donisthorpe, 1927b: 199 (Britain); Menozzi & Russo, 1930: 172 (Dominican Republic); Arnol'di, 1933b: 600 (Russia); Menozzi, 1933b: 90 (Israel genera); Karavaiev, 1936: 164 (Ukraine); Smith, M.R. 1937: 861 (Puerto Rico); Stitz, 1939: 209 (Germany); Kratochvíl, 1941: 94 (Central Europe); Novák & Sadil, 1941: 94 (Central Europe); Cole, 1942: 370 (U.S.A., Utah); Smith, M.R. 1943f: 309 (U.S.A. males); Buren, 1944a: 290 (U.S.A., Iowa); Smith, M.R. 1947f: 592 (U.S.A. genera); Creighton, 1950a: 330 (Nearctic); Kusnezov, 1956: 27 (Argentina); Brown, 1958h: 29 (New Zealand); Kusnezov, 1959: 40 (Neotropical genera); Gregg, 1963: 341 (U.S.A., Colorado); Wheeler, G.C. & Wheeler, J. 1963: 149 (U.S.A., North Dakota); Bernard, 1967: 246 (Western Europe); Wilson & Taylor, 1967: 17 (Polynesia); Bolton, 1973a: 329 (West Africa genera); Bolton & Collingwood, 1975: 3 (Britain); van Boven, 1970b: 24 (Netherlands); Snelling, R.R. & Hunt, 1976: 90 (Chile); Tarbinsky, 1976: 118 (Kyrghyzstan); van Boven, 1977: 123 (Belgium); Kutter, 1977c: 170 (Switzerland); Arnol'di & Dlussky, 1978: 546 (former European U.S.S.R.); Collingwood, 1978: 80 (Iberian Peninsula); Collingwood, 1979: 32 (Fennoscandia and Denmark); Greenslade, 1979: 28 (South Australia genera); Schembri & Collingwood, 1981: 420 (Malta); Allred, 1982: 443 (U.S.A., Utah); Baroni Urbani, 1984: 80 (Neotropical genera); Gösswald, 1985: 314 (Germany); Collingwood, 1985: 242 (Saudi Arabia); Wheeler, G.C. & Wheeler, J. 1986g: 55 (U.S.A., Nevada); Agosti & Collingwood, 1987b: 278 (Balkans); Dlussky, et al. 1990: 168 (Turkmenistan); Kupyanskaya, 1990: 152 (Far Eastern Russia); Morisita, et al., 1991: 2 (Japan); Atanasov & Dlussky, 1992: 49 (Bulgaria); Lattke, in Jaffe, 1993: 147 (Neotropical genera); Arakelian, 1994: 70 (Armenia); Wu, J. & Wang, 1995: 115 (China genera); Kupyanskaya, 1995: 331 (Far Eastern Russia); Collingwood & Agosti, 1996: 359 (Saudi Arabia); Seifert, 1996b: 161 (Central Europe); Skinner & Allen, 1996: 40 (Britain); Collingwood & Prince, 1998: 20 (Portugal); Kim & Kim, 1999: 18 (Korea); Shattuck, 1999: 26, 64 (Australia genera, synopsis); Andersen, 2000: 59 (northern Australia genera); Zhou, 2001b: 149 (China, Guangxi); Czechowski, et al. 2002: 134 (Poland); Aktaç & Radchenko, 2002: 54 (Turkey genera); Yoshimura & Onoyama, 2002: 425 (Japan genera, males); Mackay & Mackay, 2002: 221 (U.S.A., New Mexico); Palacio & Fernández, in Fernández, 2003d: 239 (Neotropical genera); Radchenko, 2005b: 186 (North Korea); Coovert, 2005: 102 (U.S.A., Ohio); Clouse, 2007b: 190 (Micronesia); Seifert, 2007: 147 (North and Central Europe); Fisher, 2009: 51 (Malagasy genera); Terayama, 2009: 196 (Taiwan); Heterick, 2009: 28 (south-western Australia genera); Boer, 2010: 14 (Benelux); Yoshimura & Fisher, 2011: 7 (Malagasy males key); Czechowski, et al. 2012: 342 (Poland); General & Alpert, 2012: 70 (Philippines genera key) ; Dlussky & Perfilieva, 2014: 433 (British Eocene species key); Baccaro, et al. 2015: 67, 126 (Brazil genera key, text).