Myrmicinae

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Myrmicinae
Mesostruma exolympica
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Lepeletier de Saint-Fargeau, 1835
Diversity
148 genera
7,861 species
40 fossil genera
204 fossil species

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.

Identification

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.

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Keys including this Subfamily

Keys to Genus in this Subfamily

Distribution

Distribution and Species Richness based on AntMaps

Statistics

Extant Taxa

Tribes Valid Genera % World Genera Invalid Genera Valid Species/Subsp. % World Species Invalid Species/Subsp.
6 148 28.8% 264 7,861 51.64% 2,492

Fossil Taxa

Fossil Genera % World Fossil Genera Valid Fossil Species/Subsp. % World Fossil Species/Subsp.
40 23.4% 204 22.57%

Fossils known from: Aix-en-Provence, France (Late Oligocene), Arkansas amber, Malvern, Arkansas, United States (Lutetian, Middle Eocene), Baltic amber, Baltic Sea region, Europe (Bartonian, Middle to Late Eocene), Bembridge Marls, Isle of Wight, UK (Priabonian, Late Eocene), Bitterfeld amber, Baltic Sea region, Europe (Bartonian, Middle to Late Eocene), Bolshaya 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, Baltic Sea region, Europe (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

Tribes

Extant Genera

AcanthognathusAcanthomyrmexAcromyrmexAdelomyrmexAdlerziaAllomerusAmoimyrmexAncyridrisAnillomyrmaAphaenogasterApterostigmaAretidrisAtopomyrmexAttaAustromoriumBaracidrisBariamyrmaBasicerosBlepharidattaBondroitiaCalyptomyrmexCardiocondylaCarebaraCataulacusCephalotesChelanerChimaeridrisColobostrumaCrematogasterCryptomyrmexCyattaCyphoidrisCyphomyrmexDacatriaDacetinopsDacetonDiaphoromyrmaDicroaspisDilobocondylaDiplomoriumDolopomyrmexEpelysidrisEpopostrumaErromyrmaEurhopalothrixEutetramoriumFormicoxenusFormosimyrmaGaoligongidrisGauromyrmexGoniommaHarpagoxenusHuberiaHylomyrmaIndomyrmaIshakidrisKalathomyrmexKartidrisKempfidrisLachnomyrmexLasiomyrmaLenomyrmexLeptothoraxLiomyrmexLophomyrmexLordomyrmaMalagidrisManicaMayriellaMegalomyrmexMelissotarsusMeranoplusMesostrumaMessorMetaponeMicrodacetonMonomoriumMycetagroicusMycetarotesMycetomoelleriusMycetophylaxMycetosoritisMycocepurusMyrmecinaMyrmicaMyrmicariaMyrmicocryptaNesomyrmexNovomessorOchetomyrmexOctostrumaOcymyrmexOrectognathusOxyepoecusOxyopomyrmexParamycetophylaxParatopulaParatrachymyrmexPatagonomyrmexPerissomyrmexPeronomyrmexPhalacromyrmexPheidolePilotrochusPodomyrmaPoecilomyrmaPogonomyrmexPristomyrmexProattaProcryptocerusPropodilobusProtalaridrisPseudoattaRecurvidrisRhopalomastixRhopalothrixRogeriaRomblonellaRostromyrmexRotastrumaRoyidrisSecostrumaSericomyrmexSolenopsisStegomyrmexStenammaStereomyrmexStrongylognathusStrumigenysSyllophopsisTalaridrisTemnothoraxTeratanerTetheamyrmaTetramoriumTrachymyrmexTranopeltaTrichomyrmexTricytarusTropidomyrmexTyrannomyrmexVeromessorVitsikaVollenhoviaVombisidrisWasmanniaXenomyrmexXerolitor

Fossil Genera

Notes

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.

Biology

Flight Period

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Flight Months for Genera within this Subfamily:

Taxon Month
Acalama
Acanthidris
Acanthognathus
Acanthomyrmex
Acidomyrmex
Acrocoelia
Acromyrmex
Adelomyrmex
Adlerzia
Aeromyrma
Afromyrma
Afroxyidris
Agastomyrma
Alistruma
Allomerus
Allopheidole
Amauromyrmex
Amoimyrmex
Ancyridris
Aneleus
Anergates
Anergatides
Anillomyrma
Anisopheidole
Antichthonidris
Antillaemyrmex
Aphaenogaster
Apomyrmex
Apsychomyrmex
Apterocrema
Apterostigma
Aratromyrmex
Archaeomyrmex
Archeatta
Arctomyrmex
Aretidris
Arnoldidris
Asemorhoptrum
Asketogenys
Aspididris
Atopogyne
Atopomyrmex
Atopula
Atta
Attaichnus
Attomyrma
Austromorium
†Acrostigma
†Afromyrma
†Agastomyrma
†Attaichnus
†Biamomyrma
†Bilobomyrma
†Boltonidris
†Brachytarsites
†Cephalomyrmex
†Clavipetiola
†Electromyrmex
†Enneamerus
†Eocenidris
†Eocenomyrma
†Eomyrmex
†Exocryptocerus
†Fallomyrma
†Fushunomyrmex
†Heteromyrmex
†Hypopomyrmex
†Ilemomyrmex
†Lampromyrmex
†Lelejus
†Lonchomyrmex
†Miosolenopsis
†Myrmecites
†Nothomyrmica
†Orbigastrula
†Oxyidris
†Parameranoplus
†Paraphaenogaster
†Plesiomyrmex
†Proleptothorax
†Propodomyrma
†Protomyrmica
†Quadrulicapito
†Quineangulicapito
†Sinaphaenogaster
†Sinomyrmex
†Solenopsites
†Sphaerogasterites
†Stigmomyrmex
†Stiphromyrmex
†Thanacomyrmex
†Wumyrmex
†Zhangidris
Baracidris
Bariamyrma
Basiceros
Biamomyrma
Bilobomyrma
Bisolenopsis
Blepharidatta
Boltonidris
Bondroitia
Borgmeierita
Brachytarsites
Brownidris
Bruchomyrma
Brunella
Calyptomyrmex
Cardiocondyla
Cardiopheidole
Carebara
Carebarella
Carebarelloides
Cataulacus
Caulomyrma
Cephalomorium
Cephalomyrmex
Cephalotes
Cephaloxys
Cepobroticus
Ceratobasis
Ceratopheidole
Chalepoxenus
Chelaner
Chelystruma
Chimaeridris
Cladarogenys
Clarkistruma
Clavipetiola
Codiomyrmex
Codioxenus
Colobocrema
Colobostruma
Conothoracoides
Conothorax
Corynomyrmex
Crateropsis
Cratomyrmex
Creightonidris
Crematogaster
Croesomyrmex
Cryptocerus
Cryptomyrmex
Cyathocephalus
Cyathomyrmex
Cyatta
Cyphoidris
Cyphomannia
Cyphomyrmex
Dacatria
Dacetinops
Daceton
Dacetum
Dacryon
Decacrema
Decamorium
Decapheidole
Descolemyrma
Diagyne
Diaphoromyrma
Dichothorax
Dicroaspis
Dilobocondyla
Diplomorium
Diplorhoptrum
Dodecamyrmica
Dodous
Dolopomyrmex Mar
Dorisidris
Doronomyrmex
Dorothea
Dyclona
Dyomorium
Dysedrognathus
Elasmopheidole
Electromyrmex
Electropheidole
Emeryia
Eneria
Enneamerus
Eocenidris
Eocenomyrma
Eomyrmex
Epelysidris
Ephebomyrmex
Epiatta
Epimyrma
Epipheidole
Epitritus
Epixenus
Epoecus
Epopostruma
Equestrimessor
Erebomyrma
Eriopheidole
Erromyrma
Eucrema
Eucryptocerus
Euophthalma
Eurhopalothrix
Eutetramorium
Fallomyrma
Forelifidis
Forelomyrmex
Formicoxenus
Formosimyrma
Fushunomyrmex
Gallardomyrma
Gaoligongidris
Gauromyrmex
Geognomicus
Gesomyrmex
Glamyromyrmex
Glyptomyrmex
Gonepimyrma
Goniomma
Goniothorax
Granisolenopsis
Gymnomyrmex
Hagioxenus
Harnedia
Harpagoxenus
Hendecapheidole
Hendecatella
Heptacondylus
Heptastruma
Hercynia
Heteromyrmex
Hexadaceton
Holcomyrmex
Huberia
Hylidris
Hylomyrma
Hypocryptocerus
Hypopomyrmex
Icothorax
Idrisella
Ilemomyrmex
Indomyrma
Ireneella
Ireneidris
Ireneopone
Irogera
Ischnomyrmex
Ishakidris
Isolcomyrmex
Isopheidole
Janetia
Kalathomyrmex
Kartidris
Kempfidris
Kyidris
Labauchena
Labidogenys
Lachnomyrmex
Laparomyrmex
Lasiomyrma
Lecanomyrma
Lelejus
Lenomyrmex
Leonomyrma
Leptomyrma
Leptothorax
Lilidris
Limnomyrmex
Liomyrmex
Lobognathus
Lobomyrmex
Lonchomyrmex
Loncyda
Lophomyrmex
Lordomyrma
Lundella
Machomyrma
Macromischa
Macromischoides
Macropheidole
Malagidris
Manica
Martia
Mayriella
Megalomyrmex
Meia
Melissotarsus
Meranoplus
Mesocrema
Mesomyrma
Mesostruma
Messor
Metapone
Miccostruma
Microdaceton
Miosolenopsis
Mitara
Monomorium
Mycetagroicus
Mycetarotes
Mycetomoellerius
Mycetophylax
Mycetosoritis
Mychothorax
Mycocepurus
Myrafant
Myrmammophilus
Myrmecina
Myrmecinella
Myrmecites
Myrmetaerus
Myrmica
Myrmicaria
Myrmicocrypta
Myrmisaraka
Myrmoxenus
Myrmus
Nematocrema
Neoatta
Neoblepharidatta
Neocrema
Neomyrma
Neostruma
Nesomyrmex
Nimbamyrma
Nothidris
Notomyrmex
Novomessor
Nystalomyrma
Ochetomyrmex
Octella
Octostruma
Ocymyrmex
Odontomyrmex
Oecodoma
Oecophthora
Oedaleocerus
Oligomyrmex
Orbigastrula
Orectognathus
Oreomyrma
Otomyrmex
Oxyepoecus
Oxygyne
Oxyidris
Oxyopomyrmex
Paedalgus
Palaeatta
Paracrema
Paracryptocerus
Parameranoplus
Paramycetophylax
Paramyrmica
Paranamyrma
Paraphacota
Paraphaenogaster
Parapheidole
Paratopula
Paratrachymyrmex
Parholcomyrmex
Parvimyrma
Patagonomyrmex
Pentastruma
Perissomyrmex
Peronomyrmex
Phacota
Phalacromyrmex
Pharaophanes
Pheidolacanthinus
Pheidole
Pheidologeton
Phidola
Phidole
Phidologeton
Physatta
Physocrema
Pilotrochus
Planimyrma
Platystruma
Plesiomyrmex
Podomyrma
Poecilomyrma
Pogonomyrmex
Polyhomoa
Pristomyrmex
Proatta Apr
Procryptocerus
Prodicroaspis
Promeranoplus
Promyrma
Propodilobus
Proscopomyrmex
Prosopidris
Protalaridris
Protholcomyrmex
Protomognathus
Protomyrmica
Pseudoatta
Pseudopodomyrma
Pyramica
Quadristruma
Quadrulicapito
Quineangulicapito
Recurvidris
Rhachiocrema
Rhopalomastix
Rhopalothrix
Rhoptromyrmex
Rogeria
Romblonella
Rostromyrmex
Rotastruma
Royidris
Santschia
Schizopelta
Scrobopheidole
Secostruma
Sericomyrmex
Serrastruma
Sifolinia
Sinomyrmex
Smithistruma
Solenomyrma
Solenops
Solenopsis
Solenopsites
Sommimyrma
Spelaeomyrmex
Sphaerocrema
Sphaerogasterites
Sporocleptes
Stegomyrmex
Stegopheidole
Stenamma
Stenothorax
Stereomyrmex
Stigmomyrmex
Stiphromyrmex
Strongylognathus
Strumigenys
Syllophopsis
Symbiomyrma
Symmyrmica
Sympheidole
Synsolenopsis
Talaridris
Teleutomyrmex
Temnothorax
Terataner
Tetheamyrma
Tetramorium
Tetramyrma
Tetrogmus
Thanacomyrmex
Theryella
Tingimyrmex
Tomognathus
Trachymyrmex
Trachypheidole
Tranetera
Tranopelta
Tranopeltoides
Trichomyrmex
Trichoscapa
Tricytarus
Triglyphothrix
Trigonogaster
Tropidomyrmex
Tyrannomyrmex
Veromessor
Vitsika
Vollenhovenia
Vollenhovia
Vombisidris
Wasmannia
Weberidris
Weberistruma
Wessonistruma
Wheeleria
Wheeleriella
Wheelerimyrmex
Willowsiella
Wumyrmex
Xenhyboma
Xenoaphaenogaster
Xenometra
Xenomyrmex
Xerolitor
Xeromyrmex
Xiphocrema
Xiphomyrmex
Zacryptocerus
Zhangidris

Morphology

Karyotype Data

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

Known Diploid Counts: 4, 8, 12, 14, 16, 18, 20, 21, 22, 24, 26, 27, 28, 30, 32, 33, 34, 36, 37, 38, 39, 40, 41, 42, 44, 45, 46, 48, 49, 50, 51, 52, 54, 56, 58, 60, 61, 62, 66, 70.

Diploid Count Details: 12 (Taxon: Pheidole mus), 14 (Taxon: Mycetarotes carinatus), 14 (Taxon: Tetramorium lanuginosum), 14 (Taxon: Tetramorium simillimum), 14 (Taxon: Tetramorium kheperra), 14 (Taxon: Tetramorium walshi), 16 (Taxon: Strumigenys), 16 (Taxon: Meranoplus bicolor), 16 (Taxon: Dacetinops concinnus), 16 (Taxon: Monomorium sahlbergi), 16 (Taxon: Pheidole), 18 (Taxon: Temnothorax tuberum), 18 (Taxon: Pheidole soritis), 18 (Taxon: Pheidole woodmasoni), 18 (Taxon: Eurhopalothrix), 18 (Taxon: Mayriella abstinens), 18 (Taxon: Pheidole), 18 (Taxon: Temnothorax nigriceps), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole), 18 (Taxon: Pheidole tepicana), 18 (Taxon: Temnothorax unifasciatus), 18 (Taxon: Mycetomoellerius urichii), 18 (Taxon: Temnothorax congruus), 20 (Taxon: Pheidole dentata), 20 (Taxon: Pheidole desertorum), 20 (Taxon: Monomorium orientale), 20 (Taxon: Epopostruma), 20 (Taxon: Pheidole fervida), 20 (Taxon: Pheidole hyatti), 20 (Taxon: Pheidole pallidula), 20 (Taxon: Pheidole fallax), 20 (Taxon: Pheidole pallidula), 20 (Taxon: Pheidole megacephala), 20 (Taxon: Pheidole subarmata), 20 (Taxon: Pheidole indica), 20 (Taxon: Tetramorium seneb), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Basiceros convexiceps), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole), 20 (Taxon: Pheidole binghamii), 20 (Taxon: Pheidole dentigula), 20 (Taxon: Pheidole porcula), 20 (Taxon: Pheidole spininodis), 20 (Taxon: Pheidole strobeli), 20 (Taxon: Pheidole capellinii), 20 (Taxon: Pheidole plagiaria), 20 (Taxon: Pheidole hortensis), 20 (Taxon: Cyphomyrmex costatus), 20 (Taxon: Tetramorium brevidentatum), 20 (Taxon: Trachymyrmex septentrionalis), 20 (Taxon: Tetramorium pnyxis), 20 (Taxon: Mycetomoellerius holmgreni), 20 (Taxon: Mycetomoellerius iheringi), 20 (Taxon: Mycetomoellerius relictus), 21 (Taxon: Monomorium indicum), 21 (Taxon: Pheidole), 22 (Taxon: Meranoplus minor), 22 (Taxon: Monomorium indicum), 22 (Taxon: Colobostruma), 22 (Taxon: Strumigenys doriae), 22 (Taxon: Tetramorium insolens), 22 (Taxon: Monomorium pharaonis), 22 (Taxon: Amoimyrmex bruchi), 22 (Taxon: Amoimyrmex silvestrii), 22 (Taxon: Amoimyrmex striatus), 22 (Taxon: Atta bisphaerica), 22 (Taxon: Atta colombica), 22 (Taxon: Atta laevigata), 22 (Taxon: Atta robusta), 22 (Taxon: Atta sexdens), 22 (Taxon: Atta sexdens), 22 (Taxon: Atta sexdens), 22 (Taxon: Chelaner), 22 (Taxon: Tetramorium adelphon), 22 (Taxon: Monomorium rothsteini), 22 (Taxon: Monomorium viridum), 22 (Taxon: Lordomyrma), 22 (Taxon: Meranoplus), 22 (Taxon: Meranoplus), 22 (Taxon: Apterostigma steigeri), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Monomorium), 22 (Taxon: Aphaenogaster japonica), 22 (Taxon: Aphaenogaster subterranea), 22 (Taxon: Orectognathus darlingtoni), 22 (Taxon: Pheidole), 22 (Taxon: Tetramorium pacificum), 22 (Taxon: Pristomyrmex), 22 (Taxon: Pheidole germaini), 22 (Taxon: Monomorium minimum), 22 (Taxon: Colobostruma alinodis), 22 (Taxon: Solenopsis molesta), 22 (Taxon: Tetramorium guineense), 22 (Taxon: Cyphomyrmex cornutus), 22 (Taxon: Solenopsis fugax), 22 (Taxon: Solenopsis fugax), 22 (Taxon: Temnothorax nylanderi), 22 (Taxon: Orectognathus versicolor), 22 (Taxon: Formicoxenus provancheri), 22 34 (Taxon: Aphaenogaster smythiesii), 24 (Taxon: Pheidole pallidula), 24 (Taxon: Apterostigma madidiense), 24 (Taxon: Apterostigma mayri), 24 (Taxon: Strumigenys friedae), 24 (Taxon: Temnothorax longispinosus), 24 (Taxon: Strumigenys dohertyi), 24 (Taxon: Temnothorax interruptus), 24 (Taxon: Chelaner whitei), 24 (Taxon: Crematogaster), 24 (Taxon: Pristomyrmex punctatus), 24 (Taxon: Crematogaster), 24 (Taxon: Crematogaster), 24 (Taxon: Crematogaster biroi), 24 (Taxon: Crematogaster longispina), 24 (Taxon: Cyphomyrmex transversus), 24 (Taxon: Temnothorax spinosior), 26 (Taxon: Tetramorium spinosum), 26 (Taxon: Mycetophylax morschi), 26 (Taxon: Mycetophylax morschi), 26 (Taxon: Leptothorax acervorum), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Tetramorium smithi), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster), 26 (Taxon: Crematogaster nawai), 26 (Taxon: Temnothorax rugatulus), 26 (Taxon: Carebara), 27 (Taxon: Cardiocondyla minutior), 27 (Taxon: Temnothorax rugatulus), 28 (Taxon: Temnothorax mediterraneus), 28 (Taxon: Mycetophylax morschi), 28 (Taxon: Mycetophylax morschi), 28 (Taxon: Tetramorium eleates), 28 (Taxon: Myrmicocrypta), 28 (Taxon: Cardiocondyla nuda), 28 (Taxon: Pheidole), 28 (Taxon: Strongylognathus huberi), 28 (Taxon: Formicoxenus chamberlini), 28 (Taxon: Formicoxenus quebecensis), 30 (Taxon: Mycetophylax conformis), 30 (Taxon: Mycetophylax conformis), 30 (Taxon: Mycetophylax morschi), 30 (Taxon: Mycetophylax morschi), 30 (Taxon: Aphaenogaster), 30 (Taxon: Myrmicocrypta), 30 (Taxon: Myrmicocrypta), 30 (Taxon: Pheidole), 30 (Taxon: Orectognathus clarki), 30 (Taxon: Cardiocondyla minutior), 30 46 (Taxon: Aphaenogaster beccarii), 32 (Taxon: Pogonomyrmex badius), 32 (Taxon: Pogonomyrmex occidentalis), 32 (Taxon: Pogonomyrmex barbatus), 32 (Taxon: Chelaner), 32 (Taxon: Solenopsis aurea), 32 (Taxon: Pogonomyrmex montanus), 32 (Taxon: Solenopsis richteri), 32 (Taxon: Solenopsis xyloni), 32 (Taxon: Pogonomyrmex subnitidus), 32 (Taxon: Aphaenogaster osimenseis), 32 (Taxon: Wasmannia auropunctata), 32 (Taxon: Wasmannia auropunctata), 32 (Taxon: Solenopsis invicta), 32 (Taxon: Aphaenogaster senilis), 32 (Taxon: Pheidole), 32 (Taxon: Pogonomyrmex californicus), 32 (Taxon: Pogonomyrmex maricopa), 32 (Taxon: Pogonomyrmex magnacanthus), 32 (Taxon: Solenopsis saevissima), 32 (Taxon: Pogonomyrmex apache), 32 (Taxon: Proatta butteli), 32 (Taxon: Pogonomyrmex comanche), 32 (Taxon: Pogonomyrmex desertorum), 32 (Taxon: Cyphomyrmex rimosus), 32 (Taxon: Pogonomyrmex rugosus), 32 (Taxon: Pogonomyrmex brevispinosus), 32 (Taxon: Carebara), 33 (Taxon: Vollenhovia), 34 (Taxon: Leptothorax muscorum), 34 (Taxon: Leptothorax canadensis), 34 (Taxon: Monomorium subopacum), 34 (Taxon: Aphaenogaster depilis), 34 (Taxon: Aphaenogaster famelica), 34 (Taxon: Aphaenogaster gibbosa), 34 (Taxon: Aphaenogaster iberica), 34 (Taxon: Monomorium), 34 (Taxon: Aphaenogaster sardoa), 34 (Taxon: Aphaenogaster testaceopilosa), 34 (Taxon: Aphaenogaster tipuna), 34 (Taxon: Vollenhovia), 34 (Taxon: Carebara), 36 (Taxon: Mycetophylax simplex), 36 (Taxon: Mycetophylax simplex), 36 (Taxon: Leptothorax muscorum), 36 (Taxon: Leptothorax canadensis), 36 (Taxon: Acromyrmex ameliae), 36 (Taxon: Strumigenys mutica), 36 (Taxon: Vollenhovia emeryi), 36 (Taxon: Aphaenogaster fulva), 36 (Taxon: Aphaenogaster miamiana), 36 (Taxon: Pogonomyrmex huachucanus), 36 (Taxon: Pheidole), 36 (Taxon: Vollenhovia), 36 (Taxon: Crematogaster), 36 (Taxon: Crematogaster brunnea), 36 (Taxon: Crematogaster subnuda), 36 (Taxon: Carebara), 36 (Taxon: Carebara), 37 (Taxon: Pheidole nodus), 38 (Taxon: Strumigenys), 38 (Taxon: Trichomyrmex scabriceps), 38 (Taxon: Acromyrmex ambiguus), 38 (Taxon: Trichomyrmex glaber), 38 (Taxon: Acromyrmex ambiguus), 38 (Taxon: Acromyrmex aspersus), 38 (Taxon: Acromyrmex coronatus), 38 (Taxon: Acromyrmex crassispinus), 38 (Taxon: Acromyrmex crassispinus), 38 (Taxon: Acromyrmex disciger), 38 (Taxon: Acromyrmex echinatior), 38 (Taxon: Acromyrmex heyeri), 38 (Taxon: Acromyrmex hispidus), 38 (Taxon: Acromyrmex lundii), 38 (Taxon: Acromyrmex molestans), 38 (Taxon: Acromyrmex niger), 38 (Taxon: Acromyrmex nigrosetosus), 38 (Taxon: Acromyrmex rugosus), 38 (Taxon: Acromyrmex subterraneus), 38 (Taxon: Acromyrmex subterraneus), 38 (Taxon: Acromyrmex balzani), 38 (Taxon: Acromyrmex balzani), 38 (Taxon: Lophomyrmex), 38 (Taxon: Aphaenogaster lamellidens), 38 (Taxon: Pheidole), 38 (Taxon: Pheidole), 38 (Taxon: Solenopsis), 38 (Taxon: Pheidole fervens), 38 (Taxon: Pheidole nodus), 38 (Taxon: Carebara), 38 (Taxon: Lophomyrmex bedoti), 39 (Taxon: Pheidole nodus), 4 (Taxon: Strumigenys louisianae), 40 (Taxon: Strumigenys godeffroyi), 40 (Taxon: Veromessor andrei), 40 (Taxon: Harpagoxenus sublaevis), 40 (Taxon: Vollenhovia), 40 (Taxon: Crematogaster), 40 (Taxon: Strumigenys diabola), 40 41 42 44 (Taxon: Aphaenogaster rudis), 41 (Taxon: Messor), 42 (Taxon: Monomorium), 42 (Taxon: Pheidole), 42 (Taxon: Aphaenogaster treatae), 42 (Taxon: Carebara diversa), 42 (Taxon: Pheidole latinoda), 42 (Taxon: Cyphomyrmex transversus), 42 (Taxon: Carebara), 42 (Taxon: Carebara), 44 (Taxon: Messor aciculatus), 44 (Taxon: Strumigenys godeffroyi), 44 (Taxon: Leptothorax muscorum), 44 (Taxon: Leptothorax canadensis), 44 (Taxon: Myrmica scabrinodis), 44 (Taxon: Manica rubida), 44 (Taxon: Allomerus octoarticulatus), 44 (Taxon: Carebara asina), 44 (Taxon: Myrmicaria brunnea), 44 (Taxon: Cephalotes pusillus), 44 (Taxon: Carebara), 44 (Taxon: Carebara), 45 46 (Taxon: Aphaenogaster longiceps), 46 (Taxon: Myrmica sabuleti), 46 (Taxon: Myrmica rubra), 46 (Taxon: Myrmica schencki), 48 (Taxon: Myrmica rubra), 48 (Taxon: Myrmica ruginodis), 49 (Taxon: Vollenhovia), 49 (Taxon: Podomyrma adelaidae), 50 (Taxon: Sericomyrmex parvulus), 50 (Taxon: Megalomyrmex incisus), 50 (Taxon: Sericomyrmex amabilis), 50 (Taxon: Vollenhovia), 50 (Taxon: Podomyrma adelaidae), 50 (Taxon: Crematogaster rothneyi), 51 (Taxon: Podomyrma adelaidae), 52 (Taxon: Cardiocondyla wroughtonii), 54 (Taxon: Mycetarotes parallelus), 56 (Taxon: Myrmica sulcinodis), 56 (Taxon: Crematogaster), 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: Mycocepurus goeldii), 8 (Taxon: Mycocepurus goeldii), 8 (Taxon: Stenamma brevicorne).

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.

  • 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.

MYRMICINAE

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.

References