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Subfamily: Dorylinae   Leach, 1815 


Taxonomic History (provided by Barry Bolton, 2016)

Extant: 1 tribe, 18 genera, 684 species, 120 subspecies

Fossil: 1 genus, 8 species

Dorylida Leach, 1815: 147 . Type-genus: Dorylus. AntCat AntWiki HOL

Taxonomic history

Dorylinae as family: Leach, 1815: 147 [Dorylida (family-group name)]; Haliday, 1836: 331 [Dorylidae]; Swainson & Shuckard, 1840: 175 [Dorylidae]; Shuckard, 1840a PDF: 188 [Dorylidae]; Smith, 1859c PDF: 1 [Dorylidae]; Mayr, 1866b PDF: 895 [Dorylidae]; Smith, 1871b PDF: 225 [Dorylidae]; André, 1882a}: 125 [Dorylidae]; Cresson, 1887 PDF: 93 [Dorylidae]; Emery, 1894h: 381 [Dorylidae]; Ashmead, 1905c PDF: 381 [Dorylidae]; Ashmead, 1906 PDF: 21 [Dorylidae]; Bernard, 1951c: 1046 [Dorylidae]; Bernard, 1953b PDF: 217 [Dorylidae].
Dorylinae as subfamily of Dorylidae: Ashmead, 1905c PDF: 381; Ashmead, 1906 PDF: 25.
Dorylinae as subfamily of Formicidae: Mayr, 1865 PDF: 16 [Dorylidae]; Emery, 1877b PDF: 70 [Dorylidae]; Forel, 1878c PDF: 365 [Dorylidae]; Emery & Forel, 1879a: 465 [Dorylidae]; André, 1881c PDF: 64 [Dorylidae]; Forel, 1892k PDF: 220 [Dorylidae]; Forel, 1893b PDF: 163; Dalla Torre, 1893 PDF}: 1; Forel, 1895b PDF: 118 [Dorylidae]; Emery, 1895l PDF: 764 [subfamily spelled Dorylini]; Emery, 1896e PDF: 174; Forel, 1899B: 22; Forel, 1901a PDF: 462; Forel, 1901d PDF: 139; Emery, 1901b PDF: 36; Bingham, 1903 PDF: 1; Emery, 1910b PDF: 3; Wheeler, 1910a: 137; Arnold, 1915: 110; Escherich, 1917: 2 [Dorylini]; Forel, 1917 PDF: 239; Bondroit, 1918 PDF: 14 [Dorylitae]; Wheeler, 1920 PDF: 53; Wheeler, 1922: 632; Clark, 1951 PDF: 16; Brown, 1954e PDF}: 28; Borgmeier, 1955 PDF: 51; all subsequent authors.
Dorylinae as dorylomorph subfamily of Formicidae: Bolton, 2003 PDF: 36, 147; Brady & Ward, 2005 PDF: 593.
Dorylinae as formicoid subfamily of Formicidae: Moreau, Bell, et al. 2006: 102.
Dorylinae as formicoid dorylomorph subfamily of Formicidae: Brady, Schultz, et al. 2006: 18173; Ward, 2007C PDF: 555.
Tribe of Dorylinae: Dorylini.
Subfamily, tribe and genus references
[Note: from Bolton 2003; letters after publication dates may not correspond.]Shuckard, 1840: 195 (genera key); Roger, 1863b: 20, 41, 42 (Typhlopone, Anomma, Dorylus, Rhogmus,Dichthadia catalogues); Smith, F. 1858b: 110, 112 (Typhlopone, Anomma diagnoses); Mayr, 1863: 394, 407,408, 453, 457 (Anomma, Dichthadia, Dorylus, Rhogmus, Typhlopone catalogues); Mayr, 1865: 16, 17(Dorylinae, Typhlopone, Anomma, Dorylus, Rhogmus, Dichthadia diagnoses); Mayr, 1867a: 91 (Typhloponediagnosis); Fore!, 1878: 365 (diagnosis); Andre, 1882c: 251 (Europe & Algeria species key); Dalla Torre,1893: 8, 9 (Rhogmus, Anomma, Dory ius) catalogues); Emery, 1895e: 699, 701, 706; 764 (diagosis genus,subgenera, all species key; diagnoses subfamily & tribe); Emery, 1896b: 174 (genera key); Fore!, 1901a: 462(subgenera key); Bingham, 1903: 2 (diagnosis, India, Sri Lanka & Burma species key); Ashmead, 1906: 22,25, 26 (subfamilies, tribes & genera keys); Wheeler, W.M. 1910d: 137 (diagnosis); Emery, 1910b: 4(diagnosis, tribes, key, catalogue); Emery, 1910b: 8 (D. (Dichthadia) diagnosis, catalogue); Emery, 1910b: 9(D. (Dorylus) diagnosis, catalogue); Emery, 1910b: 10 (D. (Anomma) diagnosis, catalogue); Emery, 1910b:12 (D. (Typhlopone) diagnosis, catalogue); Emery, 1910b: 13 (D. (Rhogmus) diagnosis, catalogue); Emery,1910b: 14 (D. (Alaopone) diagnosis, catalogue); Santschi, 1912: 154 (D. (Anomma) species key); Arnold,1915: 113, 114, 115 (tribes key, subgenera key, South Africa D. (Dorylus) species key); Fore!, 1917: 239(synoptic classification); Santschi, 1919a: 231 (D. (Rhogmus) males key); Fore!, 1921: 135 (diagnosis);Wheeler, W.M. 1922a: 39, 41, 633, 727 (subfamily & genus diagnoses, tribes & subgenera key, Afrotropicalcatalogue); Morley, 1939: 114 (phylogeny); Santschi, 1939a: 152 (D. (Alaopone) species, males key); Brown& Nutting, 1950: 123 (venation & phylogeny); Chapman & Capco, 1951: 9 (Asia checklist); Brown, 1954b:28 (phylogeny); Raignier & Boven, 1955: 86, 114 (D. (Anomma) species key); Borgmeier, 1955: 57 (tribeskey); Eisner, 1957: 478 (proventriculus morphology); Wilson, 1964: 436 (Indo-Australian species key);Gotwald, 1969: 49 (mouthparts morphology); Wheeler, G.C. & Wheeler, J. 1972: 36 (diagnosis); Kempf,1972a: 265 (Neotropical, synoptic classification); Boven, 1972: 142 (D. (Anomma) queens key; Brown,1973b: 166 (genera & distribution); Boven, 1975: 196 (D. (Dorylus) queens key; Wheeler, G.C. & Wheeler,J. 1976: 46 (larvae, review & synthesis); Smith, D.R. 1979: 1326 (North America catalogue); Gotwald &Burdette, 1981: 78 (phylogeny); Gotwald, 1982: 168 (Dorylus subgenera key); Snelling, 1981: 390 (synoptic classification); Wheeler, G.C. & Wheeler, J. 1985: 256 (synoptic classification); Billen, 1986: 168 (Dufour'sgland); Dlussky & Fedoseeva, 1988: 79 (synoptic classification); Holldobler & Wilson, 1990: 9 onward(synoptic classification, world genera, keys); Bolton, 1990c: 1357 (diagnosis, morphology, phylogeny);Baroni Urbani, Bolton & Ward, 1992: 316 (phylogeny); Xu, 1994a: 118 (China species key); Bolton, 1994:35 (diagnosis, synoptic classification); Bolton, 1995a: 1049 (census); Bolton, 1995b: 177 (catalogue);Holldobler, Obermayer & Peeters, 1996: 158 (metatibial gland); Brady, Fisher, Schultz & Ward, 2014 PDF (phylogeny); Boudinot, 2015 PDF: 48 (male diagnosis)


Afrotropical Region: Angola, Benin, Botswana, Cameroon, Central African Republic, Chad, Congo, Democratic Republic of Congo, Equatorial Guinea, Eritrea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Kenya, Lesotho, Liberia, Malawi, Mali, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, South Africa, Sudan, Swaziland, Tanzania, Togo, Uganda, Zambia, Zimbabwe
Australasia Region: Australia, Barrow Island, New Caledonia, New Guinea, Solomon Islands, Vanuatu
Indomalaya Region: Andaman and Nicobar Islands, Arunachal Pradesh, Bangladesh, Borneo, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Nicobar Island, Philippines, Singapore, Sri Lanka, Taiwan, Thailand, Vietnam
Malagasy Region: Comoros, Madagascar, Mauritius, Mayotte, Reunion, Seychelles
Nearctic Region: Alabama, Arizona, Arkansas, California, Colorado, Florida, Georgia, Illinois, Louisiana, Mississippi, Missouri, Nevada, New Mexico, North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Utah
Neotropical Region: Argentina, Belize, Bolivia, Brazil, Chile, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Galapagos Islands, Grenada, Guadeloupe, Guatemala, Guyana, Honduras, Jamaica, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Suriname, Trinidad and Tobago, Turks and Caicos Islands, Uruguay, Venezuela
Oceania Region: Fiji, Guam, Hawaii, Marshall Islands, Micronesia, Northern Mariana Islands, Samoa
Palearctic Region: Afghanistan, Algeria, Armenia, Balearic Islands, China, Egypt, Greece, Iran, Israel, Italy, Japan, Lebanon, Macaronesia, Morocco, Oman, Saudi Arabia, Tunisia, Turkey, Turkmenistan, United Arab Emirates, Yemen

Taxonomic Treatment (provided by Plazi)

Treatment Citation: Wheeler, W. M., 1922, The ants collected by the American Museum Congo Expedition., Bulletin of the American Museum of Natural History 45, pp. 39-269


Worker and Soldier.-Clypeus as a rule very short and not limited by sutures. Frontal carinae vertical, not covering the insertions of the antennae. Antennae inserted near the mouth and close to each other, often less than 12-jointed. Palpi at most 3-jointed, in LeptanillaHNS only one-jointed. Ocelli and eyes often absent (without exception in all African genera). Sutures of the thorax more or less vestigial; mesonotum touching the epinotom on the dorsal face, without interposed metanotum. Spurs of the tibiae pectinate or rudimentary. Postpetiole not always separated by a constriction from the third segment; however, in EcitonHNS, AenictusHNS, and LeptanillaHNS, narrowed into the second joint of a two-jointed pedicel. Sting developed.

Female.-Permanently apterous, with the abdomen much enlarged and swollen,' very different morphologically from the worker. Clypeus as in the worker. Frontal carinae more or less separated. Antennae 10-to 12-jointed. No ocelli; eyes not more developed than in the worker; female blind when the worker is so. Segmentation of the thorax more or less rudimentary; no traces of wings or a rudiment left at the tegulae ( DorylusHNS). Postpetiole never separated from the third segment, the pedicel always composed of one segment. Gaster long and voluminous.

Male.-Clypeus and frontal carinae much as in the female. Mandibles developed, as a rule large; in LeptanillaHNS very short. Antennae 13-jointed; scape long, in LeptanillaHNS only slightly longer than the second joint. Eyes and ocelli well developed. Thorax with normal segmentation, winged. Postpetiole and pedicel much as in the female. Genitalia completely retractile (Dorylini and Ecitini) or exserted and not retractile (Leptanillini); subgenital lamina split or furcate; cerci absent.

larvae more or less cylindrical, with short hairs, without hooked setae; mandibles small, slender, falcate.

Nymphs usually naked; enclosed in a cocoon in some species of EcitonHNS.

The three castes in this subfamily are so different from one another that their true relations remained for a very long time unsettled. The winged males were the first to be known and were originally placed with the Mutillidae. The workers and females were recognized as ants but at first classified in genera by themselves. Though their relations were more or less suspected by Lepeletier de Saint-Fargeau, Haliday, and Shuckard, the true affinities of the male and worker became only gradually known after 1850, when Savage observed for the first time in West Africa DorylusHNS males walking in an army of AnommaHNS workers. The females, leading a permanently subterranean life, are still excessively rare in collections and known only for a few species; their capture in the smaller species is rather fortuitous, whereas in such fierce army ants as AnommaHNS it is a very troublesome operation.

G. Arnoldi gives the following general account of the habits of this subfamily:

The members of this subfamily are commonly known as driver or legionary ants. The males, which are winged and provided with eyes, are frequently taken at lights; on the other hand, the workers are blind, with the exception of some species of EcitonHNS, in which there is a pair of single-faceted eyes, and the females (excepting one species of EcitonHNS) are both blind and wingless. The members of the genus DorylusHNS are almost entirely subterranean in their mode of life, rarely coming to the surface except in dull, cloudy weather. The species of the subgenus AnommaHNS, which live in the more tropical and forested regions of Africa, and to which the term driver ants was originally applied, and the Ecitini of South America, are, however, usually seen above the surface, although, should the rays of the sun prove too powerful, they will construct temporarily tunnels with particles of earth held together by their saliva. The species of AenictusHNS are not so shy of the light and may be seen foraging about even in bright sunlight. It is probable that all, or at least the majority of the species are carnivorous, although D. orientalisHNS has been shown by Green to feed also on tubers and the bark of trees.

As far as known the members of this subfamily do not as a rule make permanent nests. This course is determined by their exceedingly predatory habits, which compel the adoption of a migratory form of life together with the formation of temporary nests in localities which are sufficiently productive of animal life to detain them for any length of time. Ranging far and wide in search of prey, which consists of any animal they are strong enough to overpower, these ants must sooner or later exhaust the areas round their nests, and are forced to remove the latter to new and more productive hunting grounds.

But little is known of the habits of the Leptanillini; all species are hypogaeic. Santschi found the nest of Leptanilla nana SantschiHNS 40 cm. beneath the surface in clay soil; he caught females and workers by inundating the soil so as to force them to come out of their burrows; workers have also been taken by sifting decayed leaves. The males are attracted by lights.

A detailed account of the migrations and habits of some of the African species is given below (see under Dorylus bequaertiHNS, D. opacusHNS, D. kohliHNS, D. nigricansHNS, D. wilverthiHNS, and D. fulvusHNS).

The Dorylinae are abundantly found in all tropical parts of the world, with the exception of the Antilles and the Malagasy Region; they are absent from the larger part of Australia. A few species reach North Africa, the coasts of Asia Minor, and the central and southern United States.

1 1915, Ann. South African Mus., XIV, p. 110.

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