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Mesostruma exolympica
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Lepeletier de Saint-Fargeau, 1835
148 genera
7,785 species
40 fossil genera
193 fossil species
Evolutionary Placement



Amblyoponinae - see relationships



Ponerinae - see relationships



Dorylinae - see relationships


Dolichoderinae - see relationships

Myrmeciinae - see relationships within Myrmecia

Pseudomyrmecinae - see relationships

Formicinae - see relationships

Ectatomminae - see relationships

Myrmicinae - see relationships

See Phylogeny of Formicidae for details.

This is the largest subfamily in Australia, based on both the number of genera and the number of species. Myrmicines range greatly in size, with the smallest about 1 mm long and the largest up to 10 mm. While many species are generalist predators, some specialise on selected soft-bodied invertebrates such as Collembola and others are important seed harvesters. Workers can be found foraging at all times of the day and night, sometimes in large numbers. Nests can be found in almost any suitable location from deep in the soil to the upper branches of trees. Colonies are generally small with a few hundred to a few thousand workers, although some species can have huge nests with many thousands of workers while others form very small nests with fewer than 50 individuals.

Morphologically, these ants are very diverse. Many groups are highly modified with unusual mandibles, elongate spines, elaborate hairs or unique structures not seen in any other ants. Because of this, many genera are relatively easy to identify as they have highly distinctive features which are easily seen (given the appropriate magnification). At the same time, some genera are much less specialised or modified, and separating these from close relatives can be difficult. In these cases, close inspection of subtle differences may be required.

Myrmicines occur throughout the world in all major habitats (except arctic and antarctic regions). They are the largest subfamily of ants with over 6700 species and subspecies and 155 genera. Australia has over 350 species placed in 38 genera. Eight of the genera are restricted to Australia.


The mesosoma is attached to the gaster by two distinct segments, the petiole and postpetiole. The mandibles are generally triangular, but if they are elongate then there are no teeth along the inner margin and they are attached near the middle of the front margin of the head. The eyes are almost always present, conspicuous and with many facets but are absent in a few species. The frontal lobes are always present and expanded towards the sides so they cover the inner part of the antennal bases where they are inserted into the head when viewed from the front. The pronotum and mesonotum are fused into a single plate.

Species of myrmicines are most likely to be confused with species of Leptanilla or Tetraponera because of the two segmented petiole. However, both Leptanilla and Tetraponera have the pronotum and mesonotum unfused and with a flexible joint between them, while in all myrmicines these two plates are fused into a single structure.

Boudinot (2015) - Male Myrmicinae are uniquely identified by the strongly petiolated third abdominal segment (postpetiole), axial helcium, 1,1 maximum ventroapical tibial spur count, unvaulted abdominal tergum IV, and presence of propodeal lobes. All myrmicines lack jugal lobes and have posteriorly-situated antennal toruli, but are highly variable otherwise: mandibles fully-developed to nub-like; antenna 8–13-merous; forewing with (0)1–8 eight closed cells; and petiole sessile to long-pedunculate. Some myrmicines, e.g., Adelomyrmex and Acanthognathus, have extremely reduced wing venation similar to Leptanillinae; all myrmicine taxa examined during this study with reduced wing venation have conspicuous propodeal lobes, differentiating them easily from Leptanillinae despite secondary petiolation of abdominal segment III in some leptanillines.

AntWeb icon 02.png See images of genera within this subfamily

Keys including this Subfamily

Keys to Genus in this Subfamily


Distribution and Species Richness based on AntMaps


Extant Taxa

Tribes Valid Genera % World Genera Invalid Genera Valid Species/Subsp. % World Species Invalid Species/Subsp.
6 148 28.9% 264 7,785 51.8% 2,473

Fossil Taxa

Fossil Genera % World Fossil Genera Valid Fossil Species/Subsp. % World Fossil Species/Subsp.
40 24.2% 193 22.24%

Fossils known from: Aix-en-Provence, France (Late Oligocene), Arkansas amber, Malvern, Arkansas, United States (Lutetian, Middle Eocene), Baltic amber (Bartonian, Middle to Late Eocene), Bembridge Marls, Isle of Wight, UK (Priabonian, Late Eocene), Bitterfeld amber (Bartonian, Middle to Late Eocene), Bol’shaya Svetlovodnaya, Sikhote-Alin, Russia (Priabonian, Late Eocene), Borneo amber (Miocene), Brunn-Vösendorf, Austria (Late Miocene), Brunstatt, Haut-Rhin, France (Early Oligocene), Danish-Scandinavian amber (Bartonian, Middle to Late Eocene), Dominican amber, Dominican Republic (Burdigalian, Early Miocene), Epecuen Formation, Salinas de Hidalgo, La Pampa Prov., Argentina (Huayquerian, Late Miocene), Ethiopian amber (Early Miocene), Florissant, Colorado, United States (Late Eocene), Fushun amber, Liaoning, China (Ypresian, Early Eocene), Green River Formation, Colorado, United States (Lutetian, Middle Eocene), Chôjaburu, Iki Island, Japan (Middle Miocene), Khronya Cape, Kerch, Crimea, Russian Federation (Sarmatian, Upper Miocene), Kishenehn Formation shale, Montana, United States (Lutetian, Middle Eocene), Kleinkems, Germany (Early Oligocene), Mexican amber, Chiapas, Mexico (Middle Miocene), Mokrina (Krottensee), Czech Republic (Late Burdigalian, Early Miocene), Oeningen, Switzerland (Messinian, Late Miocene), Orapa kimberlitic deposits, Botswana (Turonian, Late Cretaceous), Parschlug, Austria (Serravallian, Miocene), Quesnel, British Columbia, Canada (Early Miocene?), Radoboj, Croatia (Burdigalian, Early Miocene), Rott, Westphalia, Germany (Late Oligocene), Rovno amber (Priabonian, Late Eocene), Sakhalin amber, Ukraine (Thanetian, Paleocene), Schossnitz (= Sosnica?), Silesia, Poland (Late Miocene), Shanwang, China (Early Miocene), Sicilian amber, Italy (Late/Upper Miocene), Vishnevaya Balka Creek, Stavropol, Russian Federation (Middle Miocene).

List of Tribes and Genera



Extant Genera


Fossil Genera



Boudinot (2015) - Of all the ant subfamilies, the Myrmicinae will be the grand challenge to understand with respect to males. At the time of writing, 139 valid genera and 6,500 valid species are described. Males of at least 30 genera are unknown, but as generic delimitation is still very active in the Myrmicinae some uncertainty exists for this number. Based on a study of the New World genera (B. Boudinot, in prep.), distinctions between genera may be weak and in many cases genera will have to be keyed multiple times due to variability. The recent subfamily-wide phylogeny of Ward et al. (2015) will contribute significantly to improving the classification of the Myrmicinae.


Karyotype Data

Explore Karyotype Data: All, Drilldown
Click here to show/hide karyotype count data.

Known Haploid Counts: 4, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 28, 30.

Haploid Count Details: 10 (Taxon: Temnothorax algerianus), 10 (Taxon: Temnothorax bernardi), 10 (Taxon: Pheidole pallidula), 10 (Taxon: Pheidole pallidula), 10 (Taxon: Pheidole fallax), 10 (Taxon: Trachymyrmex septentrionalis), 10 (Taxon: Pheidole dentigula), 10 (Taxon: Meranoplus), 10 (Taxon: Epopostruma), 10 (Taxon: Temnothorax adlerzi), 10 (Taxon: Temnothorax ravouxi), 10 (Taxon: Temnothorax corsicus), 10 (Taxon: Temnothorax gordiagini), 10 (Taxon: Temnothorax kraussei), 10 (Taxon: Temnothorax andrei), 10 (Taxon: Pheidole), 10 (Taxon: Pheidole), 10 (Taxon: Pheidole), 10 (Taxon: Temnothorax stumperi), 10 (Taxon: Pheidole), 10 (Taxon: Mycetomoellerius relictus), 10 (Taxon: Mycetomoellerius holmgreni), 10 (Taxon: Mycetomoellerius iheringi), 10 (Taxon: Tetramorium brevidentatum), 10 (Taxon: Pheidole), 10 (Taxon: Pheidole fervida), 11 (Taxon: Temnothorax nylanderi), 11 (Taxon: Temnothorax nylanderi), 11 (Taxon: Monomorium), 11 (Taxon: Monomorium), 11 (Taxon: Monomorium), 11 (Taxon: Solenopsis molesta), 11 (Taxon: Formicoxenus provancheri), 11 (Taxon: Solenopsis), 11 (Taxon: Amoimyrmex striatus), 11 (Taxon: Temnothorax sordidulus), 11 (Taxon: Acanthomyrmex), 11 (Taxon: Acanthomyrmex), 11 (Taxon: Temnothorax exilis), 11 (Taxon: Solenopsis fugax), 11 (Taxon: Aphaenogaster smythiesii), 11 (Taxon: Monomorium pharaonis), 11 (Taxon: Atta colombica), 11 (Taxon: Orectognathus darlingtoni), 11 (Taxon: Orectognathus versicolor), 11 (Taxon: Temnothorax americanus), 11 (Taxon: Leptothorax gredleri), 11 (Taxon: Lordomyrma), 11 (Taxon: Temnothorax rottenbergii), 11 (Taxon: Temnothorax flavicornis), 11 (Taxon: Colobostruma alinodis), 11 (Taxon: Aphaenogaster subterranea), 11 (Taxon: Vollenhovia), 11 16 17 (Taxon: Aphaenogaster gibbosa), 12 (Taxon: Temnothorax muellerianus), 12 (Taxon: Temnothorax corticalis), 12 (Taxon: Temnothorax spinosior), 12 (Taxon: Leptothorax), 12 (Taxon: Leptothorax), 12 (Taxon: Crematogaster), 12 (Taxon: Strumigenys dohertyi), 12 (Taxon: Temnothorax), 12 (Taxon: Temnothorax interruptus), 12 (Taxon: Temnothorax recedens), 12 (Taxon: Temnothorax kutteri), 12 (Taxon: Temnothorax longispinosus), 12 (Taxon: Temnothorax longispinosus), 12 (Taxon: Pristomyrmex punctatus), 12 (Taxon: Cyphomyrmex transversus), 13 (Taxon: Leptothorax), 13 (Taxon: Mycetophylax morschi), 13 (Taxon: Leptothorax sphagnicola), 13 (Taxon: Temnothorax melas), 13 (Taxon: Strumigenys), 13 (Taxon: Leptothorax acervorum), 13 (Taxon: Tetramorium spinosum), 14 (Taxon: Tetramorium forte), 14 (Taxon: Temnothorax parvulus), 14 (Taxon: Leptothorax), 14 (Taxon: Tetramorium caespitum), 14 (Taxon: Tetramorium caespitum), 14 (Taxon: Tetramorium semilaeve), 14 (Taxon: Tetramorium caespitum), 14 (Taxon: Temnothorax rugatulus), 14 (Taxon: Strongylognathus huberi), 14 (Taxon: Myrmecina graminicola), 14 (Taxon: Temnothorax), 14 (Taxon: Myrmecina americana), 14 (Taxon: Temnothorax mediterraneus), 14 (Taxon: Pristomyrmex), 14 (Taxon: Temnothorax lichtensteini), 15 (Taxon: Leptothorax), 15 (Taxon: Formicoxenus nitidulus), 15 (Taxon: Mycetophylax conformis), 15 (Taxon: Mycetophylax morschi), 15 (Taxon: Leptothorax faberi), 16 (Taxon: Leptothorax), 16 (Taxon: Pogonomyrmex barbatus), 16 (Taxon: Leptothorax), 16 (Taxon: Solenopsis geminata), 16 (Taxon: Solenopsis geminata), 16 (Taxon: Solenopsis invicta), 16 (Taxon: Solenopsis aurea), 16 (Taxon: Pogonomyrmex subnitidus), 16 (Taxon: Solenopsis xyloni), 16 (Taxon: Aphaenogaster osimenseis), 16 (Taxon: Pogonomyrmex maricopa), 16 (Taxon: Leptothorax canadensis), 16 (Taxon: Leptothorax faberi), 16 (Taxon: Solenopsis saevissima), 16 (Taxon: Temnothorax spinosus), 16 (Taxon: Pheidole), 16 (Taxon: Carebara), 16 (Taxon: Solenopsis richteri), 16 (Taxon: Aphaenogaster senilis), 16 (Taxon: Pogonomyrmex comanche), 17 (Taxon: Leptothorax), 17 (Taxon: Leptothorax), 17 (Taxon: Aphaenogaster famelica), 17 (Taxon: Aphaenogaster iberica), 17 (Taxon: Temnothorax exilis), 17 (Taxon: Leptothorax muscorum), 17 (Taxon: Leptothorax canadensis), 17 (Taxon: Leptothorax crassipilis), 17 (Taxon: Temnothorax gredosi), 17 (Taxon: Leptothorax retractus), 17 (Taxon: Pheidole fervens), 17 (Taxon: Pheidole), 17 (Taxon: Aphaenogaster testaceopilosa), 17 (Taxon: Pheidole nodus), 18 (Taxon: Temnothorax niger), 18 (Taxon: Harpagoxenus canadensis), 18 (Taxon: Leptothorax muscorum), 18 (Taxon: Mycetophylax simplex), 18 (Taxon: Leptothorax canadensis), 18 (Taxon: Leptothorax crassipilis), 18 (Taxon: Leptothorax pocahontas), 18 (Taxon: Leptothorax retractus), 18 (Taxon: Pheidole fervens), 18 (Taxon: Carebara sauteri), 18 (Taxon: Pheidole), 18 (Taxon: Crematogaster subnuda), 18 (Taxon: Vollenhovia emeryi), 18 (Taxon: Pheidole nodus), 18 20 (Taxon: Aphaenogaster rudis), 19 (Taxon: Acromyrmex coronatus), 19 (Taxon: Pheidole fervens), 19 (Taxon: Pheidole fervens), 19 (Taxon: Pheidole nodus), 20 (Taxon: Pheidole nodus), 20 (Taxon: Harpagoxenus sublaevis), 20 (Taxon: Stenamma westwoodii), 20 (Taxon: Pheidole fervens), 21 (Taxon: Messor barbarus), 21 (Taxon: Leptothorax), 21 (Taxon: Temnothorax luteus), 21 (Taxon: Temnothorax racovitzai), 22 (Taxon: Leptothorax), 22 (Taxon: Myrmica scabrinodis), 22 (Taxon: Messor aciculatus), 22 (Taxon: Leptothorax muscorum), 22 (Taxon: Temnothorax ambiguus), 22 (Taxon: Leptothorax canadensis), 22 (Taxon: Cephalotes pusillus), 23 (Taxon: Myrmica rubra), 23 (Taxon: Myrmica schencki), 23 (Taxon: Temnothorax curvispinosus), 23 (Taxon: Apterostigma madidiense), 23 (Taxon: Leptothorax canadensis), 23 (Taxon: Temnothorax duloticus), 23 (Taxon: Myrmica sabuleti), 23 (Taxon: Myrmicaria), 23 (Taxon: Temnothorax angustulus), 23 24 25 (Taxon: Leptothorax kutteri), 24 (Taxon: Myrmica lobicornis), 24 (Taxon: Myrmica sulcinodis), 24 (Taxon: Temnothorax duloticus), 24 (Taxon: Myrmica ruginodis), 25 (Taxon: Temnothorax duloticus), 25 (Taxon: Megalomyrmex incisus), 25 (Taxon: Sericomyrmex), 26 (Taxon: Leptothorax pacis), 28 (Taxon: Myrmica sulcinodis), 28 (Taxon: Leptothorax goesswaldi), 30 (Taxon: Pogonomyrmex imberbiculus), 4 (Taxon: Stenamma brevicorne), 8 (Taxon: Temnothorax albipennis), 8 (Taxon: Temnothorax albipennis), 8 (Taxon: Temnothorax rougeti), 9 (Taxon: Temnothorax aveli), 9 (Taxon: Temnothorax corticalis), 9 (Taxon: Temnothorax congruus), 9 (Taxon: Temnothorax nigriceps), 9 (Taxon: Temnothorax affinis), 9 (Taxon: Temnothorax tuberum), 9 (Taxon: Temnothorax tuberum), 9 (Taxon: Temnothorax schaumii), 9 (Taxon: Temnothorax), 9 (Taxon: Mycetomoellerius), 9 (Taxon: Pheidole), 9 (Taxon: Pheidole), 9 (Taxon: Pheidole), 9 (Taxon: Mycetomoellerius urichii), 9 (Taxon: Temnothorax unifasciatus), 9 (Taxon: Temnothorax unifasciatus).

Known Diploid Counts: 4, 8, 12, 14, 16, 18, 20, 21, 22, 24, 26, 27, 28, 30, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 44, 45, 46, 48, 49, 50, 51, 52, 56, 58, 60, 61, 62, 66, 70.

Diploid Count Details: 12 (Taxon: Pheidole mus), 14 (Taxon: Tetramorium walshi), 14 (Taxon: Tetramorium lanuginosum), 14 (Taxon: Tetramorium simillimum), 14 (Taxon: Tetramorium kheperra), 14 (Taxon: Tetramorium), 16 (Taxon: Meranoplus bicolor), 16 (Taxon: Pheidole), 16 (Taxon: Monomorium sahlbergi), 16 (Taxon: Strumigenys), 16 (Taxon: Dacetinops concinnus), 18 (Taxon: Temnothorax congruus), 18 (Taxon: Temnothorax nigriceps), 18 (Taxon: Temnothorax tuberum), 18 (Taxon: Pheidole soritis), 18 (Taxon: Pheidole tepicana), 18 (Taxon: Eurhopalothrix), 18 (Taxon: Mycetomoellerius), 18 (Taxon: Tetramorium), 18 (Taxon: Tetramorium), 18 (Taxon: Tetramorium), 18 (Taxon: Mayriella abstinens), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole woodmasoni), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Mycetomoellerius urichii), 18 (Taxon: Temnothorax unifasciatus), 20 (Taxon: Pheidole hyatti), 20 (Taxon: Monomorium orientale), 20 (Taxon: Pheidole pallidula), 20 (Taxon: Pheidole pallidula), 20 (Taxon: Pheidole binghamii), 20 (Taxon: Pheidole hortensis), 20 (Taxon: Pheidole fallax), 20 (Taxon: Pheidole plagiaria), 20 (Taxon: Pheidole dentata), 20 (Taxon: Trachymyrmex septentrionalis), 20 (Taxon: Pheidole dentigula), 20 (Taxon: Cyphomyrmex costatus), 20 (Taxon: Epopostruma), 20 (Taxon: Tetramorium), 20 (Taxon: Tetramorium), 20 (Taxon: Pheidole indica), 20 (Taxon: Tetramorium), 20 (Taxon: Tetramorium), 20 (Taxon: Pheidole capellinii), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole porcula), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole spininodis), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole strobeli), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Basiceros convexiceps), 20 (Taxon: Pheidole), 20 (Taxon: Mycetomoellerius relictus), 20 (Taxon: Pheidole), 20 (Taxon: Mycetomoellerius holmgreni), 20 (Taxon: Mycetomoellerius iheringi), 20 (Taxon: Tetramorium brevidentatum), 20 (Taxon: Pheidole), 20 (Taxon: Tetramorium pnyxis), 20 (Taxon: Pheidole desertorum), 20 (Taxon: Tetramorium seneb), 20 (Taxon: Pheidole megacephala), 20 (Taxon: Pheidole subarmata), 20 (Taxon: Pheidole fervida), 20 24 32 (Taxon: Apterostigma), 21 (Taxon: Monomorium indicum), 21 (Taxon: Pheidole), 22 (Taxon: Temnothorax nylanderi), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium rothsteini), 22 (Taxon: Tetramorium pacificum), 22 (Taxon: Solenopsis molesta), 22 (Taxon: Formicoxenus provancheri), 22 (Taxon: Meranoplus), 22 (Taxon: Meranoplus), 22 (Taxon: Amoimyrmex striatus), 22 (Taxon: Amoimyrmex striatus), 22 (Taxon: Strumigenys doriae), 22 (Taxon: Acanthomyrmex), 22 (Taxon: Acanthomyrmex), 22 (Taxon: Aphaenogaster japonica), 22 (Taxon: Colobostruma), 22 (Taxon: Solenopsis fugax), 22 (Taxon: Solenopsis fugax), 22 (Taxon: Tetramorium insolens), 22 (Taxon: Monomorium pharaonis), 22 (Taxon: Apterostigma steigeri), 22 (Taxon: Atta bisphaerica), 22 (Taxon: Atta colombica), 22 (Taxon: Atta laevigata), 22 (Taxon: Atta robusta), 22 (Taxon: Orectognathus darlingtoni), 22 (Taxon: Orectognathus versicolor), 22 (Taxon: Tetramorium), 22 (Taxon: Chelaner), 22 (Taxon: Monomorium minimum), 22 (Taxon: Pheidole germaini), 22 (Taxon: Lordomyrma), 22 (Taxon: Meranoplus minor), 22 (Taxon: Pristomyrmex), 22 (Taxon: Monomorium indicum), 22 (Taxon: Pheidole), 22 (Taxon: Monomorium viridum), 22 (Taxon: Tetramorium adelphon), 22 (Taxon: Colobostruma alinodis), 22 (Taxon: Tetramorium guineense), 22 (Taxon: Aphaenogaster subterranea), 22 (Taxon: Atta sexdens), 22 (Taxon: Atta sexdens), 22 (Taxon: Atta sexdens), 22 (Taxon: Cyphomyrmex cornutus), 22 34 (Taxon: Aphaenogaster smythiesii), 24 (Taxon: Chelaner whitei), 24 (Taxon: Temnothorax spinosior), 24 (Taxon: Recurvidris), 24 (Taxon: Pheidole pallidula), 24 (Taxon: Crematogaster), 24 (Taxon: Crematogaster), 24 (Taxon: Crematogaster), 24 (Taxon: Crematogaster longispina), 24 (Taxon: Strumigenys dohertyi), 24 (Taxon: Strumigenys friedae), 24 (Taxon: Acanthomyrmex), 24 (Taxon: Temnothorax interruptus), 24 (Taxon: Apterostigma mayri), 24 (Taxon: Tetramorium), 24 (Taxon: Temnothorax longispinosus), 24 (Taxon: Pristomyrmex punctatus), 24 (Taxon: Crematogaster biroi), 24 (Taxon: Cyphomyrmex transversus), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster nawai), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Temnothorax rugatulus), 26 (Taxon: Mycetophylax morschi), 26 (Taxon: Mycetophylax morschi), 26 (Taxon: Tetramorium), 26 (Taxon: Carebara), 26 (Taxon: Leptothorax acervorum), 26 (Taxon: Tetramorium spinosum), 27 (Taxon: Temnothorax rugatulus), 27 (Taxon: Cardiocondyla minutior), 28 (Taxon: Cardiocondyla nuda), 28 (Taxon: Tetramorium caespitum), 28 (Taxon: Tetramorium caespitum), 28 (Taxon: Formicoxenus chamberlini), 28 (Taxon: Formicoxenus quebecensis), 28 (Taxon: Tetramorium eleates), 28 (Taxon: Strongylognathus huberi), 28 (Taxon: Allomerus decemarticulatus), 28 (Taxon: Mycetophylax morschi), 28 (Taxon: Mycetophylax morschi), 28 (Taxon: Temnothorax mediterraneus), 28 (Taxon: Pheidole), 30 (Taxon: Aphaenogaster), 30 (Taxon: Mycetophylax conformis), 30 (Taxon: Mycetophylax conformis), 30 (Taxon: Orectognathus clarki), 30 (Taxon: Mycetophylax morschi), 30 (Taxon: Mycetophylax morschi), 30 (Taxon: Myrmicocrypta), 30 (Taxon: Myrmicocrypta), 30 (Taxon: Pheidole), 30 46 (Taxon: Aphaenogaster beccarii), 32 (Taxon: Pogonomyrmex barbatus), 32 (Taxon: Pogonomyrmex rugosus), 32 (Taxon: Solenopsis geminata), 32 (Taxon: Solenopsis geminata), 32 (Taxon: Solenopsis geminata), 32 (Taxon: Solenopsis invicta), 32 (Taxon: Solenopsis aurea), 32 (Taxon: Pogonomyrmex montanus), 32 (Taxon: Pogonomyrmex subnitidus), 32 (Taxon: Solenopsis xyloni), 32 (Taxon: Proatta butteli), 32 (Taxon: Aphaenogaster osimenseis), 32 (Taxon: Cyphomyrmex rimosus), 32 (Taxon: Pogonomyrmex badius), 32 (Taxon: Pogonomyrmex maricopa), 32 (Taxon: Chelaner), 32 (Taxon: Pogonomyrmex californicus), 32 (Taxon: Solenopsis saevissima), 32 (Taxon: Pogonomyrmex magnacanthus), 32 (Taxon: Pheidole), 32 (Taxon: Carebara), 32 (Taxon: Solenopsis richteri), 32 (Taxon: Aphaenogaster senilis), 32 (Taxon: Pogonomyrmex apache), 32 (Taxon: Pogonomyrmex brevispinosus), 32 (Taxon: Pogonomyrmex comanche), 32 (Taxon: Pogonomyrmex desertorum), 32 (Taxon: Pogonomyrmex occidentalis), 32 (Taxon: Wasmannia auropunctata), 32 (Taxon: Wasmannia auropunctata), 33 (Taxon: Vollenhovia), 34 (Taxon: Monomorium), 34 (Taxon: Aphaenogaster sardoa), 34 (Taxon: Aphaenogaster depilis), 34 (Taxon: Aphaenogaster famelica), 34 (Taxon: Aphaenogaster gibbosa), 34 (Taxon: Aphaenogaster iberica), 34 (Taxon: Leptothorax muscorum), 34 (Taxon: Aphaenogaster tipuna), 34 (Taxon: Leptothorax canadensis), 34 (Taxon: Carebara), 34 (Taxon: Monomorium subopacum), 34 (Taxon: Aphaenogaster testaceopilosa), 34 (Taxon: Vollenhovia), 35 (Taxon: Tetramorium), 36 (Taxon: Acromyrmex ameliae), 36 (Taxon: Crematogaster), 36 (Taxon: Aphaenogaster fulva), 36 (Taxon: Aphaenogaster miamiana), 36 (Taxon: Leptothorax muscorum), 36 (Taxon: Mycetophylax simplex), 36 (Taxon: Mycetophylax simplex), 36 (Taxon: Tetramorium), 36 (Taxon: Leptothorax canadensis), 36 (Taxon: Carebara), 36 (Taxon: Carebara), 36 (Taxon: Crematogaster brunnea), 36 (Taxon: Pheidole), 36 (Taxon: Pogonomyrmex huachucanus), 36 (Taxon: Crematogaster subnuda), 36 (Taxon: Strumigenys mutica), 36 (Taxon: Vollenhovia), 36 (Taxon: Vollenhovia emeryi), 37 (Taxon: Pheidole nodus), 38 (Taxon: Acromyrmex balzani), 38 (Taxon: Acromyrmex balzani), 38 (Taxon: Acromyrmex rugosus), 38 (Taxon: Acromyrmex coronatus), 38 (Taxon: Acromyrmex disciger), 38 (Taxon: Acromyrmex molestans), 38 (Taxon: Lophomyrmex bedoti), 38 (Taxon: Acromyrmex ambiguus), 38 (Taxon: Acromyrmex ambiguus), 38 (Taxon: Acromyrmex aspersus), 38 (Taxon: Acromyrmex crassispinus), 38 (Taxon: Acromyrmex crassispinus), 38 (Taxon: Acromyrmex echinatior), 38 (Taxon: Acromyrmex heyeri), 38 (Taxon: Acromyrmex hispidus), 38 (Taxon: Acromyrmex niger), 38 (Taxon: Acromyrmex nigrosetosus), 38 (Taxon: Solenopsis), 38 (Taxon: Aphaenogaster lamellidens), 38 (Taxon: Lophomyrmex), 38 (Taxon: Acromyrmex lundii), 38 (Taxon: Pheidole fervens), 38 (Taxon: Carebara), 38 (Taxon: Pheidole), 38 (Taxon: Pheidole), 38 (Taxon: Strumigenys), 38 (Taxon: Acromyrmex subterraneus), 38 (Taxon: Trichomyrmex glaber), 38 (Taxon: Acromyrmex subterraneus), 38 (Taxon: Trichomyrmex scabriceps), 38 (Taxon: Pheidole nodus), 39 (Taxon: Pheidole nodus), 4 (Taxon: Strumigenys louisianae), 40 (Taxon: Strumigenys godeffroyi), 40 (Taxon: Harpagoxenus sublaevis), 40 (Taxon: Cardiocondyla), 40 (Taxon: Crematogaster), 40 (Taxon: Strumigenys diabola), 40 (Taxon: Veromessor andrei), 40 (Taxon: Vollenhovia), 40 41 42 44 (Taxon: Aphaenogaster rudis), 41 (Taxon: Messor), 42 (Taxon: Monomorium), 42 (Taxon: Carebara diversa), 42 (Taxon: Aphaenogaster treatae), 42 (Taxon: Pheidole latinoda), 42 (Taxon: Pheidole), 42 (Taxon: Carebara), 42 (Taxon: Carebara), 44 (Taxon: Manica rubida), 44 (Taxon: Myrmica scabrinodis), 44 (Taxon: Myrmicaria brunnea), 44 (Taxon: Allomerus octoarticulatus), 44 (Taxon: Messor aciculatus), 44 (Taxon: Leptothorax muscorum), 44 (Taxon: Leptothorax canadensis), 44 (Taxon: Myrmicaria), 44 (Taxon: Myrmicaria), 44 (Taxon: Myrmicaria), 44 (Taxon: Carebara), 44 (Taxon: Carebara), 44 (Taxon: Carebara asina), 44 (Taxon: Cephalotes pusillus), 44 (Taxon: Strumigenys godeffroyi), 45 46 (Taxon: Aphaenogaster longiceps), 46 (Taxon: Myrmica rubra), 46 (Taxon: Myrmica schencki), 46 (Taxon: Myrmica sabuleti), 46 (Taxon: Myrmicaria), 48 (Taxon: Myrmica rubra), 48 (Taxon: Myrmica ruginodis), 49 (Taxon: Podomyrma adelaidae), 49 (Taxon: Vollenhovia), 50 (Taxon: Crematogaster rothneyi), 50 (Taxon: Podomyrma adelaidae), 50 (Taxon: Megalomyrmex incisus), 50 (Taxon: Sericomyrmex), 50 (Taxon: Sericomyrmex amabilis), 50 (Taxon: Vollenhovia), 51 (Taxon: Podomyrma adelaidae), 52 (Taxon: Cardiocondyla wroughtonii), 56 (Taxon: Crematogaster), 56 (Taxon: Myrmica sulcinodis), 58 (Taxon: Pogonomyrmex imberbiculus), 60 (Taxon: Pogonomyrmex imberbiculus), 61 (Taxon: Pogonomyrmex imberbiculus), 62 (Taxon: Pogonomyrmex imberbiculus), 66 (Taxon: Myrmecina), 70 (Taxon: Erromyrma latinodis), 8 (Taxon: Stenamma brevicorne), 8 (Taxon: Mycocepurus), 8 (Taxon: Mycocepurus goeldii).


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

  • MYRMICINAE [subfamily of Formicidae]
    • Myrmicites Lepeletier de Saint-Fargeau, 1835: 169. Type-genus: Myrmica Latreille, 1804: 179.

Taxonomic History

  • Myrmicinae as group name: Lepeletier de Saint-Fargeau, 1835: 169 [Myrmicites]; Nylander, 1846a: 877 [Myrmicae].
  • Myrmicinae as family: Smith, F. 1851: 4 [Myrmicidae]; Smith, F. 1861b: 45 [Myrmicidae]; Smith, F. 1871a: 324 [Myrmicidae]; André, 1882a: 125 [Myrmicidae]; Cresson, 1887: 93 [Myrmicidae]; Emery, 1894g: 383 [Myrmicidae]; Saunders, 1896: 18 [Myrmicidae]; Ashmead, 1905b: 383 [Myrmicidae]; Novák & Sadil, 1941: 71 [Myrmicidae]; Bernard, 1951: 1058 [Myrmicidae]; Bernard, 1953b: 222 [Myrmicidae].
  • Myrmicinae as subfamily of Poneridae: Smith, F. 1858b: 114 [Myrmicidae].
  • Myrmicinae as tribe of Formicidae: André, 1874: 167 [Myrmicidae].
  • Myrmicinae as subfamily of Myrmicidae: Ashmead, 1905b: 383.
  • Myrmicinae as subfamily of Formicidae: Mayr, 1855: 290, 299 [Myrmicidae]; Smith, F. 1857a: 70 [Myrmicidae]; Mayr, 1861: 21 [Myrmicidae]; Smith, F. 1862b: 33 [Myrmicidae]; Mayr, 1862: 738 [Myrmicidae]; Mayr, 1865: 17 [Myrmicidae]; Mayr, 1868b: 24 [Myrmicidae]; Forel, 1870: 307 [Myrmicidae]; Forel, 1874: 22 [Myrmicidae]; Emery, 1877a: 70 [Myrmicidae]; Forel, 1878: 367 [Myrmicidae]; Emery & Forel, 1879a: 456 [Myrmicidae]; André, 1881b: 64 [Myrmicidae]; Nasonov, 1889: 28 [Myrmicidae]; Forel, 1891b: 11 [Myrmicidae]; Forel, 1892j: 220 [Myrmicidae]; Forel, 1893a: 163 [Myrmicinae]; Dalla Torre, 1893: 53; Emery, 1895j: 768 [subfamily spelled Myrmicini]; Emery, 1896e: 179; Forel, 1899c: 30; Forel, 1902f: 520; Bingham, 1903: 105; Ruzsky, 1905b: 103; Wheeler, W.M. 1910g: 138; Emery, 1914a: 29; Wheeler, 1915g: 806 [Myrmicides]; Wheeler, W.M. 1915h: 40; Donisthorpe, 1915d: 74; Arnold, 1916: 166; Escherich, 1917: 2 [Myrmicini]; Forel, 1917: 240 [subfamily spelled Myrmicini]; Bondroit, 1918: 14 [Myrmicitae]; Wheeler, W.M. 1920: 53; Wheeler, W.M. 1922a: 124; Emery, 1921f: 3; Karavaiev, 1934: 59; Clark, 1951: 16; Brown, 1954e: 28; Wheeler, G.C. & Wheeler, J. 1972a: 40; Brown, 1973b: 166; subsequent authors.
  • Myrmicinae as myrmicomorph subfamily of Formicidae: Bolton, 2003: 52, 182.
  • Myrmicinae as formicoid subfamily of Formicidae: Moreau, et al. 2006: 102; Brady, et al. 2006: 18173.
  • Myrmicinae as formicoid myrmicomorph subfamily of Formicidae: Ward, 2007a: 556.

Taxonomic References

Mayr, 1865: 17 (diagnosis); Mayr, 1867a: 91 (diagnosis); Forel, 1878: 367 (diagnosis); Handlirsch, 1907: 872 (*fossil taxa catalogue); Dalla Torre, 1893: 53 (catalogue); Emery, 1895j: 768 (diagnosis); Emery, 1896e: 179 (genera key); Wheeler, W.M. 1910g: 138 (diagnosis); Emery, 1912b: 101 (phylogeny); Emery, 1914a: 34 (phylogeny, tribe key); Arnold, 1916: 164 (diagnosis); Forel, 1917: 240 (synoptic classification); Forel, 1921c: 139 (diagnosis); Emery, 1921f: 3 (diagnosis, tribes and genera key, catalogue); Wheeler, W.M. 1922a: 124, 655 (diagnosis, tribes key); Brown & Nutting, 1950: 126 (venation, phylogeny); Brown, 1954e: 28 (phylogeny); Eisner, 1957: 477 (proventriculus morphology); Bernard, 1967: 93 (diagnosis); Gotwald, 1969: 99 (mouthparts morphology); Wheeler, G.C. & Wheeler, J. 1972a: 40 (diagnosis); Brown, 1973b: 166 (genera, distribution); Wheeler, G.C. & Wheeler, J. 1976b: 52 (larvae, review and synthesis); Kugler, C. 1978a: 413 (sting structure); Kugler, C. 1978b: 267 (pygidial glands); Kugler, C. 1979c: 117 (sting, evolution); Snelling, R.R. 1981: 393 (synoptic classification); Caetano, F.H. 1984: 257 (digestive tract, morphology); Wheeler, G.C. & Wheeler, J. 1985: 257 (synoptic classification); Billen, 1986: 167 (Dufour's gland); Dlussky & Fedoseeva, 1988: 79 (synoptic classification); Hölldobler & Wilson, 1990: 9 onward (synoptic classification, genera keys); Baroni Urbani, et al. 1992: 317 (phylogeny); Bolton, 1994: 75 (diagnosis, synoptic classification, genera keys); Bolton, 1995a: 1040 (census); Bolton, 1995b: 13 (catalogue); Hashimoto, 1996: 354 (phylogenetic position); Baroni Urbani, 2000: 480 (phylogeny); Dlussky & Rasnitsyn, 2002: 422 (diagnosis for wingless fossils); Bolton, 2003: 52, 182 (diagnosis, synopsis); Moreau, et al. 2006: 102 (phylogeny); Brady, et al. 2006: 18173 (phylogeny); Ward, 2007a: 556 (classification); Keller, 2011: 1 (morphology, phylogeny); Ward, et al. 2015: 61 (phylogeny); Boudinot, 2015: 56 (diagnosis); Fisher & Bolton, 2016: 50 (diagnosis).

Regional and National Faunas with Keys

Mayr, 1855: 391 (Austria); Mayr, 1861: 29 (Europe); Mayr, 1868b: 79 (*Baltic Amber); André, 1874: 171 (Europe); Forel, 1874: 29 (Switzerland); Saunders, E. 1880: 213 (Britain); André, 1882c: 256 (Europe and Algeria); Cresson, 1887: 98 (U.S.A. genera); Provancher, 1887: 243 (Canada); Nasonov, 1889: 54 (Russia); Forel, 1891b: 11 (Madagascar genera); Lameere, 1892: 66 (Belgium); Forel, 1902f: 520 (India and Sri Lanka genera); Bingham, 1903: 105 (India, Sri Lanka and Burma); Ruzsky, 1905b: 103 (Russian Empire); Wasmann, 1906: 13 (Luxemburg); Bondroit, 1910: 490 (Belgium); Wheeler, W.M. 1910g: 558 (North America genera); Stitz, 1914: 55 (Central Europe); Gallardo, 1915: 32 (Argentina genera); Forel, 1915d: 8 (Switzerland); Donisthorpe, 1915d: 74 (Britain); Arnold, 1916: 166, 170 (South Africa tribes, genera); Emery, 1916b: 112 (Italy); Wheeler, W.M. 1916m: 581 (U.S.A., Connecticut); Bondroit, 1918: 90 (France and Belgium); Kutter, 1920b: 144 (Switzerland); Soudek, 1922: 20 (Czechoslovakia); Stärcke, 1926: 84 (Netherlands); Karavaiev, 1927c: 256 (Ukraine); Donisthorpe, 1927b: 77 (Britain); Menozzi & Russo, 1930: 170 (Dominican Republic); Gallardo, 1932b: 91 (Argentina, tribes); Arnol'di, 1933b: 596 (Russia); Menozzi, 1933b: 88 (Israel genera); Karavaiev, 1934: 60 (Ukraine); Smith, M.R. 1937: 829 (Puerto Rico); Stitz, 1939: 63 (Germany); Kratochvíl, 1941: 71 (Central Europe); Novák & Sadil, 1941: 71 (Central Europe); Cole, 1942: 360 (U.S.A., Utah); Smith, M.R. 1943f: 291 (U.S.A., males); Holgersen, 1943b: 166 (Norway); Holgersen, 1944: 198 (Norway); Buren, 1944a: 281 (U.S.A., Iowa); Smith, M.R. 1947f: 543 (U.S.A. genera); van Boven, 1947: 170 (Belgium); Creighton, 1950a: 83 (Nearctic); Kusnezov, 1956: 15 (Argentina); Brown, 1958h: 25 (New Zealand); van Boven, 1959: 7 (Netherlands); Gregg, 1963: 288 (U.S.A., Colorado); Wheeler, G.C. & Wheeler, J. 1963: 92 (U.S.A., North Dakota); Collingwood, 1964: 94 (Britain); Bernard, 1967: 95 (Western Europe); Wilson & Taylor, 1967: 13 (Polynesia); van Boven, 1970b: 9 (Netherlands); Kempf, 1972a: 263 (Neotropical, synoptic classification); Bolton, 1973a: 325 (West Africa genera); Bolton & Collingwood, 1975: 3 (Britain); Snelling, R.R. & Hunt, 1976: 70 (Chile); Tarbinsky, 1976: 19 (Kyrghyzstan); van Boven, 1977: 69 (Belgium); Kutter, 1977c: 31 (Switzerland); Arnol'di & Dlussky, 1978: 524 (former European U.S.S.R.); Collingwood, 1978: 75 (Iberian Peninsula); Collingwood, 1979: 36 (Fennoscandia and Denmark); Greenslade, 1979: 20 (South Australia genera); Schembri & Collingwood, 1981: 423 (Malta); Allred, 1982: 438 (U.S.A., Utah); Baroni Urbani, 1984: 76 (Neotropical genera); Verhaeghe, Deligne, et al. 1984: 112 (Belgium genera); Gösswald, 1985: 289 (Germany); Collingwood, 1985: 245 (Saudi Arabia); Wheeler, G.C. & Wheeler, J. 1986g: 20 (U.S.A., Nevada); Nilsson & Douwes, 1987: 57 (Norway); Agosti & Collingwood, 1987b: 265 (Balkans); Dlussky, et al. 1990: 181 (Turkmenistan); Kupyanskaya, 1990: 89 (Far Eastern Russia); Ogata, 1991b: 61 (Japan genera); Morisita, et al. 1992: 1 (Japan); Atanasov & Dlussky, 1992: 51 (Bulgaria); Lattke, in Jaffe, 1993: 153 (Neotropical genera); Arakelian, 1994: 15 (Armenia); Wu, J. & Wang, 1995: 57 (China genera); Kupyanskaya, 1995: 327 (Far Eastern Russia); Collingwood & Agosti, 1996: 308 (Saudi Arabia); Seifert, 1996b: 108 (Central Europe); Skinner & Allen, 1996: 43 (Britain); Collingwood & Prince, 1998: 10 (Portugal); Shattuck, 1999: 39, 122 (Australia genera, synopsis); Andersen, 2000: 37 (northern Australia genera); Zhou, 2001b: 69 (China, Guangxi); Czechowski, et al. 2002: 135 (Poland); Aktaç & Radchenko, 2002: 55 (Turkey genera); Yoshimura & Onoyama, 2002: 424 (Japan genera, males); Mackay & Mackay, 2002: 58 (U.S.A., New Mexico); Palacio & Fernández, in Fernández, 2003d: 244 (Neotropical genera and synopsis); Coovert, 2005: 32 (U.S.A., Ohio); Radchenko, 2005b: 184 (North Korea); Clouse, 2007b: 190 (Micronesia); Seifert, 2007: 110 (North and Central Europe); Terayama, 2009: 131 (Taiwan); Heterick, 2009: 36 (south-western Australia genera); Boer, 2010: 46 (Benelux); Eguchi, et al. 2011: 8 (Vietnam genera); Czechowski, et al. 2012: 344 (Poland); General & Alpert, 2012: 73 (Philippines genera key) ; Dlussky & Perfilieva, 2014: 433 (British Eocene species key); Baccaro, et al. 2015: 82, 202 (Brazil genera key, text).

Taxonomic Notes

Bolton 2003:

The myrmicomorph subfamilies

Subfamilies Agroecomyrmecinae, Myrmicinae.

Diagnosis Paraglossae present on labium (note 1). Antennal sockets vertical or strongly inclined upward toward midline of head (note 2). Torulus not completely fused to frontal lobe. Ocelli absent (note 3). Promesonotal suture usually absent, less commonly vestigial; in the latter case the suture is fully fused and immobile, the pronotum and mesonotum incapable of movement relative to each other (note 4). Metacoxal cavities fully closed, the annulus broad and without a suture (note 5). Propodeal lobes usually present (note 6). Waist of two segments petiole plus postpetiole) (note 7). Petiole with complete tergosternal fusion (note 8). Presclerites present on abdominal segment IV (first gastral) (also in male) the presternite distinctly shorter than the pretergite (note 9). Pretarsal claws without a preapical tooth on the inner margin (note 10). Sting present, usually functional. Jugal lobe absent from hindwing of alates. Pupae naked. [Synopsis, p. 181.]

Notes (1) Paraglossae have not been recorded in Metapone; the presence of paraglossae is regarded as plesiomorphic. (2) For distribution of character see notes under formicomorph subfamilies. In most myrmicomorphs the antennal sockets are relatively widely separated. They are secondarily approximated in tribes such as Stenammini, Solenopsidini and Melissotarsini and secondarily extremely widely separated in Agroecomyrmecini and Cataulacini. (3) Ocelli are considered absent in myrmicomorph workers although in worker polymorphic taxa, especially those with major workers that develop gyne-like characters (for example some Pheidole, Pheidologeton, Oligomyrmex, Atta, Solenopsis geminata group, some Crematogaster depressa group (list is not exhaustive) the largest may have at least a median ocellus present. Elsewhere ocelli, or more usually the median ocellus alone, may be sporadically and perhaps teratologically developed in individuals or nests (observed in Atopomyrmex, Cataulacus, Huberia, Metapone, Monomorium). Ocelli are universal in alate queens and males. (4) In some myrmicomorph taxa a line or feeble indentation across the dorsal alitrunk may indicate the original track of the pro meso notal suture. Elsewhere in the family the suture is also univerally fused and immobile in all dorylomorph subfamilies except Leptanilloidinae, and fused in the poneromorph groups Ectatommini, Paraponerini, Proceratiini and Probolomyrmecini. Plesiomorphic lack of promesonotal fusion, with retained mobility of pronotum and mesonotum relative to one another, is characteristic of the formicomorphs, myrmeciomorphs and leptanillomorphs, as well as the dorylomorph subfamily Leptanilloidinae and all poneromorphs except those mentioned above. (5) For distribution of character see notes under Dolichoderinae. (6) Propodeallobes are universal in myrmicomorphs except for Melissotarsini and some Crematogastrini, where their absence is regarded as a secondary adaptation. For distribution of this character see notes under myrmeciomorph subfamilies. (7) All myrmicomorphs have a two-segmented waist and the character probably represents a single evolutionary event. A single-segmented waist, the plesiomorphic condition, is universal in formicomorphs and poneromorphs except for Paraponerini and a few species of Proceratiini, where segment III is quite reduced and could be termed sub-postpetiolate. Among the myrmeciomorphs, dorylomorphs and leptanillomorphs both counts occur in each group and sometimes both counts occur within a single subfamily, as is the case in Myrmeciinae, Cerapachyinae and Ecitoninae. It is obvious that the reduction of abdominal segment III to a distinct postpetiole has evolved independently many times in Formicidae. (8) For distribution of character see notes under formicomorph subfamilies. (9) The relatively short presternite on abdominal segment IV is also found in Pseudomyrmecinae. This feature has almost certainly arisen independently and is not a myrmicine/pseudomyrmecine synapomorphy. (10) Pretarsal claws are also simple, lacking a preapical tooth, in the formicomorph and leptanillomorph subfamilies. In other subfamily groups such teeth are variously developed, being absent in some taxa, present in others; the presence of such teeth is plesiomorphic.


Diagnosis. With characters of myrmicomorph subfamilies. Clypeus usually inserted between antennal sockets (note 1). Antennal sockets usually not strongly migrated laterally (note 2). Metapleural gland orifice a longitudinal slit or narrow crescent that opens dorsally to posterodorsally, not overhung by a cuticular flange or flap (note 3). Petiole in posterior view with the fused tergile and sternile equally convex, their inner margins forming a circle. Tergite and sternite of helcium together form a rough circle in frontal view, the apices of the two sclerites meet end to end, the tergile does not overlap the sternite; helcium sternite bulges ventrally and is not retracted (note 4) (also in male). Abdominal segment III (postpetiole) usually without tergosternal fusion (note 5); abdominal segment IV without tergosternal fusion (note 6), with or without a stridulitrum on pretergite. Sternite of abdominal segment IV not reduced, the segment not strongly downcurved. Postpygidial glands absent (note 7). [Synopsis, p. 182.]

Notes. (1) Median portion of clypeus fails to extend back between the antennal sockets only in Melissotarsini; this is considered to be an independent apomorphic development of that tribe. (2) The only genus in which the antennal sockets and frontal lobes are strongly migrated laterally is Cataulacus, in which feature it resembles Tatuidris (see above), but otherwise their cephalic morphologies are extremely different. (3) Vertically directed guard-hairs that arise some distance below the orifice are frequently developed. External appearance of the metapleural gland is very similar to that encountered in Ectatomminae; the structure may be synapomorphic. The slit-like orifice of the gland is usually so narrow in Myrmicinae that it is inconspicuous. (4) A ventrally bulging helcium sternite is also developed throughout the dorylomorph subfamilies, in Discothyrea (Proceratiini) and in Tatuidris (Agroecomyrmecini), but in all of these the sternite is retracted, so that tergite overlaps sternite, and the latter is attached some distance up the inner surface of the tergite. (5) Tergosternal fusion of abdominal segment III occurs only in Cataulacini, Cephalotini and Myrmicaria among all the Myrmicinae. This may be a synapomorphy in the first two but is certainly independently derived in the last. For distribution of this character elsewhere in the family see notes under dorylomorph subfamilies. (6) In Ankylomyrma the tergite of abdominal segment IV is enormously hypertrophied; the sternite appears to be a small sclerite fused to the anteroventral rim of the tergite: see under Ankylomyrmini. (7) Postpygidial glands are also absent in the formicomorph subfamilies, almost certainly independently.

Comments (i) This subfamily, overwhelmingly the largest, most diverse and most successful of the Formicidae, is divided into a number of tribes, which are gathered here into formal or informal tribe groups. Some groups of tribes are demonstrably monophyletic (the dacetine-, cephalotine-, and attine tribe groups) but the remainder are grouped only by inclusive diagnoses, so their monophyly remains in doubt. Within this latter set of groups some individual tribes are demonstrably monophyletic but others are not. A number of new, mainly monogeneric, tribes have been set up to accommodate the more obvious problem areas. In general these are taxa with a wealth of autapomorphic developments but a dearth of recognisable synapomorphies. (ii) Many of the extreme inquilines (workerless permanent social parasites) encountered in the Myrmicinae will fail taxon diagnosis because of their "inquiline syndrome" habitus and grossly convergent morphological reductions and modifications, acquired in response to their unique way of life. Inclusion of such forms in higher taxa must rest upon molecular analysis, for instance the proof by Sanetra & Buschinger (2000) that the extreme inquilines Anergates and Teleutomyrmex are correctly grouped with Tetramorium. (iii) For another possible Ectatommini/Myrmicinae synapomorphy (as well as (3) above) see comments under the former.