Dorylinae

The subfamily revision by Marek Borowiec (2016) has drastically altered the taxonomy of the constituent genera. This work created a much sounder foundation for future studies of Doryline ants. Borowiec's summary of the subfamily "The subfamily is a monophyletic group of predatory ants, occurring throughout most of the tropical and subtropical regions of the world, with an appreciable number of species in warm temperate environments. The relatively few dorylines for which foraging biology is known usually prey on other ants or social insects, although notable exceptions occur and several of the charismatic ‘army ants’ evolved more generalized predatory habits. There are about 680 described species with an estimate of the total diversity being at least 1,000. The diversity of both habits and morphology within the subfamily is high and nesting can be subterranean or arboreal, with colony sizes ranging from a few dozen to millions of workers. These workers vary from having well-developed compound eyes to being entirely blind, having very short to very long slender appendages, and with the cuticle varying from coarsely sculptured to polished and shiny, with dull or conspicuous coloration. We know about the biology of a few conspicuous species within the subfamily. Overall, our biologically knowledge is very poor because many species are subterranean or occur at low abundances. Comparative studies of doryline biology have been thwarted by poor taxonomic knowledge, a lack of identification resources, and a classification that does not reflect evolutionary relationships."

Identification
Males: Boudinot (2015) - Most Dorylinae are uniquely identified by the bidentate or pronged ninth abdominal sternum, lack of pygostyles, and poorly developed clypeus. Males of the Leptanilloides genus group are highly derived and are identifiable by the following combination of characters: antennal toruli abutting or very nearly abutting anterior clypeal margin; oblique mesopleural sulcus absent; four closed cells present on forewing; cinctus between abdominal pre- and postsclerites IV absent.

Genus richness
Genus richness by country based on regional taxon lists (countries with darker colours are more genus-rich).



Species richness
Species richness by country based on regional taxon lists (countries with darker colours are more species-rich).



Nomenclature
DORYLINAE [subfamily of Formicidae]
 * Dorylida Leach, 1815: 147 [as family-group name]. Type-genus: Dorylus.
 * Dorylidae: Haliday, 1836: 331 [emended spelling of suffix].
 * Dorylinae: Forel, 1893a: 163 [as subfamily of Formicidae]; Brown, 1954e: 28.
 * Dorylii: Forel, 1893a: 163 [as tribe of Dorylinae].
 * Dorylini: Ashmead, 1905b: 381 [emended spelling of suffix].

Bolton (2003) stated the following regarding the group of 6 subfamilies that are now all considered part of Dorylinae (Brady et al. 2014). It provides some guidance in regards to the features of the subfamily but a new taxonomic study is needed to clarify what is stated below.

The dorylomorph subfamilies

Subfamilies Aenictinae, Aenictogitoninae, Cerapachyinae, Dorylinae, Ecitoninae, Leptanilloidinae.

Diagnosis Prementum not visible when mouthparts closed, completely concealed behind labrum and outgrowths of the maxillae that meet medially. Clypeus reduced, usually very narrow from front to back and antennal sockets close to anterior margin of head (note 1). Antennal sockets horizontal, in plane of transverse axis of head and partially to entirely exposed. Eyes frequently very reduced or absent. Metatibial gland present, located distally on the ventral surface (note 2). Metacoxal cavities fully closed, without a suture in the broad annulus. Metapleural gland orifice concealed beneath a ventrally directed cuticular flap or flange. Petiole sessile to sub sessile. Abdominal segment III with complete tergosternal fusion (note 3), segment IV without tergosternal fusion (note 4). Sternite of helcium large, bulging ventrally and visible in profile (note 5) (also in male); sternite attached some distance up inner surface of tergite so that tergite overlaps sternite. Abdominal segment IV with strongly developed presclerites. Stridulitrum absent from pretergite of abdominal segment IV. ''Spiracles of abdominal segments V - VII shifted posteriorly on each segment, not concealed by the posterior margin of the preceding tergite and visible without distension or dissection. Pygidium modified: either large and with dorsum flattened and armed with teeth or spines, or reduced to a narrow V-shaped sclerite'' (note 6). Sting apparatus with furcula fused to base of sting or absent (note 7). Male with mandibles usually edentate except for apical tooth (note 8), generally falcate; cerci absent and genitalia completely retractile; hypopygium bidentate to biaculeate. Jugal lobe absent from hindwing of alates. [Synopsis, p. 137.]

Notes (1) A similar clypeal reduction, also with full exposure of the antennal sockets (see below), occurs in Apomyrminae, most Leptanillinae and some poneromorph groups. Reduction of the clypeus is morphoclinal in the dorylomorph Cerapachyinae, being relatively broad in Cylindromyrmecini, narrower in Acanthostichini and narrowest in Cerapachyini. In all other dorylomorph subfamilies the clypeus is so narrow that the antennal sockets are very close to, or at, the anterior margin of the head (also in male). Antennal sockets are usually fully exposed in dorylomorphs, being separated by no more than a crest or closely approximated pair of low, vertical carinae. Horizontal weak frontal lobes are rarely developed, see notes under Cerapachyinae. (2) Metatibial gland is secondarily absent in Leptanilloidinae and not visible in some Cerapachyinae and the ecitonine genus Nomamyrmex, where the gland has been overlaid with opaque cuticle. Elsewhere in the Formicidae a few species of Pachycondyla and Diacamma (both Ponerini) and many Dacetini have glands on the metatibia. It is suspected that these are not homologous but rather are the result of independent evolutions as in the former the gland is differently sited and in the latter the glands are located dorsally, on all tibiae and on the femora as well. Myopopone (Amblyoponini) has an enormous gland bulla on the metatibia but this is located posterodorsally and is also present on the mesotibia. (3) Tergosternal fusion of abdominal segment III is universally present in leptanillomorphs, dorylomorphs except for males of Ecitoninae and in all poneromorphs except for Adetomyrma; it is universally absent in myrmeciomorphs, all myrmicomorphs except for Cataulacus, Cephalotes and Myrmicaria; it is partial (basal only, near helium) in Plagiolepidini (Formicinae) and some genera of Dolichoderinae. (4) The presclerites but not the postsclerites of abdominal segment IV may be tergostemally fused in Cerapachyinae; elsewhere in dorylomorphs fusion is entirely absent. (5) The helcium stemite also bulges ventrally in the poneromorph genus Discothyrea and the agroecomyrmecine genus Tatuidris (which are probably parallelisms), and in the subfamily Myrmicinae. In the dorylomorphs and the first two of these taxa the helcium sternite is attached some distance up the inner wall of the tergite (anterior view) so that the tergite overlaps the sternite, whereas in Myrmicinae the helcium sternite is attached directly across the tergal apices. (6) A single species of the Ponerini genus Pachycondyla (Pachycondyla crassinoda) has the pygidium armed with a pair of teeth; it shows no other dorylomorph attributes so the character is certainly an autapomorphy of this one species. (7) A similar condition of the furcula is also seen in formicomorphs, some leptanillomorphs and in Myrmicinae with reduced stings. (8) Some species of Cerapachys males have up to 12 denticles on each mandible and a few Sphinctomyrmex have 2 - 3 teeth; the edentate condition is otherwise universal.

Comments (i) This group of subfamiles is emphatically monophyletic but the position of the group with respect to other formicids remains unclear. Although obviously related to the poneromorphs and leptanillomorphs the nature of the relationship has not yet been untangled. What does seem clear is that the dorylomorph group should not be regarded as the sister-group of all the poneromorphs together, but rather it has arisen from somewhere within that diverse assemblage. (ii) For the diagnostic reasons given below the various higher groups of dorylomorphs are regarded as subfamilies, rather than as subgroups of a single subfamily.