Dolichoderinae

Dolichoderinae
Dolichoderus dentatus
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Dolichoderinae Forel, 1878
Diversity
28 genera 851 species 23 fossil genera 148 fossil species
Synonyms
Hypoclineidae

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.

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.

See images of genera within this subfamily

Keys including this Subfamily

• Key to Iberian Peninsula Subfamilies
• Key to Australian Ant Subfamilies
• Key to Subfamilies of North America
• Key to Subfamilies, Males
• Key to subfamilies of the Neotropical region

Keys to Genus in this Subfamily

• Key to Australian Genera of Dolichoderinae
• Key to North American Genera of Dolichoderinae
• Key to the Dolichoderinae genera of the southwestern Australian Botanical Province
• Dolichoderinae of the southwestern Australian Botanical Province
• Key to Philippine Dolichoderinae
• Key to the genera of Greek Dolichoderinae

Distribution

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).

Statistics

Extant Taxa

Tribes	Valid Genera	% World Genera	Invalid Genera	Valid Species/Subsp.	% World Species	Invalid Species/Subsp.
6	28	5.4%	24	851	5.56%	326

Fossil Taxa

Fossil Genera	% World Fossil Genera	Valid Fossil Species/Subsp.	% World Fossil Species/Subsp.
23	13.1%	148	16.12%

Fossils known from: Aix-en-Provence, France (Late Oligocene), Arkansas amber, Malvern, Arkansas, United States (Lutetian, Middle Eocene), Baltic amber, Baltic Sea region, Europe (Priabonian, Late Eocene), Belarus amber, near Kobryn, Belarus (Eocene), Bembridge Marls, Isle of Wight, UK (Priabonian, Late Eocene), Bitterfeld amber, Baltic Sea region, Europe (Priabonian, Late Eocene), Bolshaya Svetlovodnaya, Sikhote-Alin, Russia (Priabonian, Late Eocene), Célas, Gard, France (Late Eocene), Chon-Tyz mine, Naryn Province, Kyrgyzstan (Middle Miocene), Danish-Scandinavian amber (Priabonian, Late Eocene), Dominican amber, Dominican Republic (Burdigalian, Early Miocene), Florissant, Colorado, United States (Late Eocene), Foremost Formation amber, Alberta, Canada (Campanian, Late Cretaceous), Green River Formation, Colorado, United States (Lutetian, Middle Eocene), Kishenehn Formation shale, Montana, United States (Lutetian, Middle Eocene), Kleinkems, Germany (Early Oligocene), Kuclín, near Bílina, Czechia (Late Eocene), Malyi Kamyshlak, Kerch, Crimea, Russian Federation (Middle Miocene), Oeningen, Switzerland (Messinian, Late Miocene), Parschlug, Austria (Serravallian, Miocene), Quesnel, British Columbia, Canada (Early Miocene?), Radoboj, Croatia (Burdigalian, Early Miocene), Rott, Westphalia, Germany (Late Oligocene), Rovno amber, Baltic Sea region, Europe (Priabonian, Late Eocene), Sakhalin amber, Ukraine (Thanetian, Paleocene), Shanwang, China (Early Miocene), Sicilian amber, Italy (Late/Upper Miocene).

List of Tribes and Genera

Tribes

Bothriomyrmecini • Dolichoderini • Leptomyrmecini • Miomyrmecini • Tapinomini • Zherichiniini

Extant Genera

Anillidris • Anonychomyrma • Aptinoma • Arnoldius • Axinidris • Azteca • Bothriomyrmex • Chronoxenus • Doleromyrma • Dolichoderus • Dorymyrmex • Ecphorella • Forelius • Froggattella • Gracilidris • Iridomyrmex • Leptomyrmex • Linepithema • Liometopum • Loweriella • Nebothriomyrmex • Ochetellus • Papyrius • Philidris • Ravavy • Tapinoma • Technomyrmex • Turneria

Fossil Genera

Alloiomma • Asymphylomyrmex • Chronomyrmex • Ctenobethylus • Damzenomyrmex • Dlusskyus • Elaeomyrmex • Elaphrodites • Eldermyrmex • Emplastus • Eotapinoma • Eurymyrmex • Kotshkorkia • Ktunaxia • Leptomyrmula • Miomyrmex • Petraeomyrmex • Proiridomyrmex • Protazteca • Schismiscapus • Usomyrma • Yantaromyrmex • Zherichinius

Evolutionary History

Boudinot (2015) - The Dolichoderinae is one of the major ant lineages, with over 700 described species distributed in 28 valid genera. Males are unknown for four genera (Ecphorella, Gracilidris, Loweriella, Nebothriomyrmex). One of the historically intractable problems of myrmecology has been the separation of male Dolichoderinae and Formicinae. Here, the telomeral character described by Yoshimura & Fisher (2011) is confirmed as a diagnostic synapomorphy of the subfamily on a global scale, while the antennal torulus to clypeus distance character was first described to the author’s knowledge in Czechowski et al. (2012). Several other characters were found to distinguish male Dolichoderinae and Formicinae, as indicated in the key and diagnosis. Genera of Dolichoderinae have been keyed globally by Shattuck (1992), although this key stands in need of updating. A key to the New World dolichoderine genera is in preparation (B. Boudinot, in prep.).

Radchencko (2023) - Dolichoderinae is one of the oldest evolutionary lineages of the modern ant subfamilies: its oldest members are known from the Late Cretaceous Canadian ambers (Campanian, 78–79 Ma), e. g. Eotapinoma macalpini and Chronomyrmex medicinehatensis (Dlussky, 1999; McKellar et al., 2013; see also Boudinot et al., 2022). Moreover, in Kachin amber (Myanmar the earliest Cenomanian, ca. 100 Ma), another undescribed dolichoderine genus and species were discovered (Boudinot et al., 2020, 2022).

The extant genera of Dolichoderinae were first found in the middle Eocene deposits (ca. 45 Ma) (Wilson, 1985; Dlussky & Rasnitsyn, 2002), but diversity of this subfamily considerably increased since the Late Eocene. Thus, in Late Eocene European ambers Dolichoderinae dominated in terms of the number of specimens, and were among the taxonomically most diverse subfamilies, inferior in this respect only to Myrmicinae and Formicinae. However, starting from the Oligocene their proportion began to decrease significantly (Dlussky, 2002, 2008; Dlussky & Rasnitsyn, 2002, 2007) and at present they are inferior to more subfamilies in the number of species and genera (as calculated based on Bolton, 2023).

Biology

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.

Flight Period

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Flight Months for Genera within this Subfamily:

Taxon	Month
Azteca	Jan • Feb • Mar • Apr • May • Jun • Jul • Aug • Sep • Oct • Nov • Dec
Dolichoderus	Feb • Mar • Apr • May • Jun • Jul • Aug • Sep • Oct • Nov
Dorymyrmex	Apr • May • Jun • Jul • Aug
Forelius	May • Jun • Jul • Aug
Iridomyrmex	Oct
Linepithema	May • Jun • Jul
Liometopum	Mar
Tapinoma	Mar
Technomyrmex	Sep

Morphology

Karyotype Data

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Known Haploid Counts: 5, 7, 8, 9, 11, 12, 13, 14, 16.

Haploid Count Details: 11 (Taxon: Bothriomyrmex corsicus), 11 (Taxon: Bothriomyrmex), 12 (Taxon: Leptomyrmex erythrocephalus), 13 (Taxon: Dorymyrmex bicolor), 14 (Taxon: Dolichoderus scabridus), 14 (Taxon: Ochetellus glaber), 16 (Taxon: Forelius mccooki), 5 (Taxon: Tapinoma melanocephalum), 5 (Taxon: Tapinoma), 7 (Taxon: Tapinoma indicum), 7 (Taxon: Doleromyrma), 7 (Taxon: Iridomyrmex), 8 (Taxon: Anonychomyrma itinerans), 8 (Taxon: Tapinoma sessile), 8 (Taxon: Papyrius nitidus), 8 (Taxon: Linepithema humile), 8 (Taxon: Tapinoma erraticum), 8 (Taxon: Tapinoma erraticum), 9 (Taxon: Iridomyrmex mattiroloi), 9 (Taxon: Iridomyrmex purpureus), 9 (Taxon: Iridomyrmex bicknelli), 9 (Taxon: Dorymyrmex thoracicus), 9 (Taxon: Technomyrmex albipes), 9 (Taxon: Technomyrmex albipes), 9 (Taxon: Iridomyrmex), 9 (Taxon: Tapinoma nigerrimum), 9 (Taxon: Tapinoma nigerrimum), 9 (Taxon: Tapinoma simrothi), 9 (Taxon: Tapinoma simrothi).

Known Diploid Counts: 10, 12, 14, 16, 18, 20, 22, 26, 28, 30, 32, 33, 38, 46, 48.

Diploid Count Details: 10 (Taxon: Dolichoderus lutosus), 10 (Taxon: Tapinoma indicum), 10 (Taxon: Tapinoma melanocephalum), 10 (Taxon: Tapinoma melanocephalum), 10 (Taxon: Tapinoma melanocephalum), 10 (Taxon: Tapinoma), 12 (Taxon: Doleromyrma), 14 (Taxon: Doleromyrma), 14 (Taxon: Iridomyrmex), 16 (Taxon: Anonychomyrma itinerans), 16 (Taxon: Philidris cordata), 16 (Taxon: Philidris cordata), 16 (Taxon: Tapinoma sessile), 16 (Taxon: Papyrius nitidus), 16 (Taxon: Linepithema humile), 16 (Taxon: Technomyrmex albipes), 16 (Taxon: Technomyrmex albipes), 16 (Taxon: Anonychomyrma), 16 (Taxon: Tapinoma erraticum), 16 (Taxon: Tapinoma erraticum), 18 (Taxon: Dorymyrmex pulchellus), 18 (Taxon: Iridomyrmex mattiroloi), 18 (Taxon: Iridomyrmex purpureus), 18 (Taxon: Iridomyrmex bicknelli), 18 (Taxon: Dolichoderus bidens), 18 (Taxon: Linepithema fuscum), 18 (Taxon: Linepithema fuscum), 18 (Taxon: Iridomyrmex anceps), 18 (Taxon: Technomyrmex albipes), 18 (Taxon: Dolichoderus), 18 (Taxon: Iridomyrmex), 18 (Taxon: Iridomyrmex), 18 (Taxon: Iridomyrmex), 18 (Taxon: Iridomyrmex), 18 (Taxon: Tapinoma nigerrimum), 18 (Taxon: Tapinoma simrothi), 18 (Taxon: Dorymyrmex pyramicus), 20 (Taxon: Dolichoderus voraginosus), 22 (Taxon: Arnoldius pusillus), 22 (Taxon: Dolichoderus diversus), 26 (Taxon: Dorymyrmex flavus), 26 (Taxon: Dorymyrmex bicolor), 28 (Taxon: Dolichoderus scabridus), 28 (Taxon: Ochetellus glaber), 28 (Taxon: Dolichoderus quadripunctatus), 28 (Taxon: Technomyrmex), 28 (Taxon: Technomyrmex), 28 (Taxon: Azteca trigona), 30 (Taxon: Dolichoderus thoracicus), 30 (Taxon: Technomyrmex), 30 (Taxon: Technomyrmex), 32 (Taxon: Forelius mccooki), 33 (Taxon: Dolichoderus thoracicus), 38 (Taxon: Dolichoderus decollatus), 38 (Taxon: Dolichoderus imitator), 46 (Taxon: Dolichoderus imitator), 48 (Taxon: Iridomyrmex anceps).

Phylogeny

Formicoidea

Camelomecia clade (unplaced to family) Camelomecia Camelosphecia Formicidae               Stem Formicidae Unplaced to subfamily (Baikuris, Cretomyrma, Dlusskyidris) Clade Sphecomyrmines Haidomyrmecinae Sphecomyrminae Zigrasimeciinae Clade Antennoclypeata Brownimeciinae Crown Formicidae               Leptanillinae Martialinae Amblyoponinae - see relationships Apomyrminae Proceratiinae Ponerinae - see relationships Agroecomyrmecinae Paraponerinae Dorylinae - see relationships Aneuretinae Dolichoderinae - see relationships Myrmeciinae - see relationships Pseudomyrmecinae - see relationships Formicinae - see relationships Ectatomminae - see relationships Myrmicinae - see relationships

See Phylogeny of Formicidae for details.

Nomenclature

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

• DOLICHODERINAE [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).

References

• Borowiec, M.L., Moreau, C.S., Rabeling, C. 2020. Ants: Phylogeny and Classification. In: C. Starr (ed.), Encyclopedia of Social Insects (doi:10.1007/978-3-319-90306-4_155-1).
• Boudinot, B.E. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European Journal of Taxonomy 120, 1-62 (http://dx.doi.org/10.5852/ejt.2015.120).
• Forel, A. 1878c. Études myrmécologiques en 1878 (première partie) avec l'anatomie du gésier des fourmis. Bull. Soc. Vaudoise Sci. Nat. 15:337-392.
• Radchenko, A.G. 2023. A New Ant Genus (Hymenoptera, Formicidae) from the Late Eocene Rovno Amber. Zoodiversity, 57(4), 323–336 (doi:10.15407/zoo2023.04.323).
• Shattuck, S.O. 1992a. Review of the dolichoderine ant genus Iridomyrmex Mayr with descriptions of three new genera (Hymenoptera: Formicidae). J. Aust. Entomol. Soc. 31:13-18.
• Shattuck, S.O. 1992b. Higher classification of the ant subfamilies Aneuretinae, Dolichoderinae and Formicinae (Hymenoptera: Formicidae). Syst. Entomol. 17:199-206.