Leptothorax

Leptothorax
Leptothorax acervorum
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Crematogastrini
Alliance:	Formicoxenus genus group
Genus:	Leptothorax Mayr, 1855
Type species
Formica acervorum, now Leptothorax acervorum
Diversity
20 species (Species Checklist, Species by Country) Leptothorax acervorum Specimen Label
Synonyms
Doronomyrmex Kutter, 1945 Mychothorax Ruzsky, 1904

Identification

See images of species within this genus

Keys including this Genus

• Key to Ant Genera of the Navajo Reservation
• Key to North American Genera of Myrmicinae
• Key to Vietnamese Myrmicinae Genera
• Key to the Ant Genera of New Mexico

 

Keys to Species in this Genus

• Key to Palaearctic Leptothorax and Formicoxenus

Distribution

Species by Region

Number of species within biogeographic regions, along with the total number of species for each region.

	Afrotropical Region	Australasian Region	Indo-Australian Region	Malagasy Region	Nearctic Region	Neotropical Region	Oriental Region	Palaearctic Region
Species	0	0	0	0	11	2	1	11
Total Species	2840	1735	3042	932	835	4378	1740	2862

Biology

Heinze and Gratiashvili (2015) - A phylogenetic analysis suggests that functional monogyny evolved convergently in several lineages of Leptothorax. Reproductive skew thus appears to be a labile trait. This is made particularly obvious by the existence of both functionally monogynous and polygynous populations of Leptothorax acervorum (Heinze et al. 1995; Gill et al. 2009).

Association with Other Organisms

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Species Uncertain

• An unknown species is a host for the cestode Choanotaenia crateriformis (a parasitoid) (Quevillon, 2018) (encounter mode secondary; indirect transmission; transmission outside nest).
• An unknown species is a prey for the Microdon fly Microdon mutabilis (a predator) (Quevillon, 2018).

All Associate Records for Genus

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Taxon	Relationship	Associate Type	Associate Taxon	Associate Relationship	Locality	Source	Notes
Leptothorax	host	cestode	Choanotaenia crateriformis	parasitoid		Quevillon, 2018	encounter mode secondary; indirect transmission; transmission outside nest
Leptothorax	prey	''Microdon'' fly	Microdon mutabilis	predator		Quevillon, 2018
Leptothorax acervorum	host	ant	Harpagoxenus sublaevis	slave maker		Beibl et al., 2005; de la Mora et al., 2021 (and included references)
Leptothorax acervorum	host	ant	Leptothorax goesswaldi	workerless inquiline
Leptothorax acervorum	host	ant	Leptothorax kutteri	workerless inquiline
Leptothorax acervorum	host	ant	Leptothorax pacis	workerless inquiline
Leptothorax acervorum	host	cestode	Choanotaenia unicoronata	parasite	South Tyrol, Italy; Bavaria, Germany	Buschinger, 1973; Peru et al., 1990; Laciny, 2021
Leptothorax acervorum	host	gregarine	Mattesia geminata	parasite	United States (Montana), Germany, Italy	Kleespies et al., 1997; Buschinger & Kleespies, 1999
Leptothorax acervorum	host	nematode	Mermithidae (unspecified "Mermix")	parasite	Germany	Gosswald, 1930; Laciny, 2021
Leptothorax acervorum	xenobiont	ant	Formica fusca	xenobiont	Finland	Czechowski, 2004; Kanizsai et al., 2013	In different successional series of rocky habitats, in mound of ''Formica lugubris''
Leptothorax calderoni	xenobiont	ant	Camponotus modoc	xenobiont	United States	Kanizsai et al., 2013; Wheeler, 1917	Pine forest. In log/in stump.
Leptothorax canadensis	host	ant	Harpagoxenus canadensis	slave maker		Beibl et al., 2005; de la Mora et al., 2021
Leptothorax faberi	workerless inquiline	ant	Leptothorax	host
Leptothorax goesswaldi	workerless inquiline	ant	Leptothorax acervorum	host
Leptothorax gredleri	host	ant	Harpagoxenus sublaevis	slave maker		Foitzik et al., 2003; Guillem et al., 2014; de la Mora et al., 2021
Leptothorax kutteri	host	gregarine	Mattesia geminata	parasite	Germany	Buschinger & Kleespies, 1999
Leptothorax kutteri	workerless inquiline	ant	Leptothorax acervorum	host
Leptothorax muscorum	host	ant	Harpagoxenus canadensis	slave maker	Québec, Canada	Smith, 1939
Leptothorax muscorum	host	ant	Harpagoxenus sublaevis	slave maker		Lenoir et al., 2001; Bauer et al., 2009; Brandt et al., 2007; Fischer-Blass et al., 2006; Foitzik et al., 2003; de la Mora et al., 2021
Leptothorax muscorum	host	gregarine	Mattesia geminata	parasite	United States (Montana), Germany, Canada (Alberta), Europe	Kleespies et al., 1997; Buschinger & Kleespies, 1999
Leptothorax muscorum	host	gregarine	Neogregarinorida, Ophryocystidae (species unspecified)	parasite	United States (Montana)	Buschinger et al., 1995	reported as ''Leptothorax muscorum'' complex, includes European species
Leptothorax muscorum	mutualist	aphid	Rhopalosiphum nymphaeae	trophobiont		Jones, 1927; Saddiqui et al., 2019
Leptothorax muscorum	xenobiont	ant	Formica rufa	xenobiont	Sweden	Kanizsai et al., 2013; Wheeler, 1901
Leptothorax muscorum	xenobiont	ant	Myrmica rubra	xenobiont	Finland	Czechowski, 2004; Kanizsai et al., 2013	Shore meadow. Under stone
Leptothorax pacis	workerless inquiline	ant	Leptothorax acervorum	host		Buschinger et al., 1981
Leptothorax paraxenus	workerless inquiline	ant	Leptothorax sp. B	host		Heinze & Alloway, 1992
Leptothorax wilsoni	workerless inquiline	ant	Leptothorax	host

Flight Period

All Flight Records for Genus

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Taxon	Month	Source	Notes
Leptothorax acervorum	Jun • Jul • Aug • Sep	antkeeping.info
Leptothorax gredleri	Jul • Aug	antkeeping.info
Leptothorax kutteri	Jul • Aug	antkeeping.info
Leptothorax muscorum	Jun • Jul • Aug • Sep	antkeeping.info
Leptothorax pacis	Jul	antkeeping.info

Life History Traits

• Mean colony size: 25-60 (Greer et al., 2021)
• Compound colony type: inquilinism (Greer et al., 2021)
• Nest site: hypogaeic (Greer et al., 2021)
• Diet class: omnivore (Greer et al., 2021)
• Foraging stratum: subterranean/leaf litter (Greer et al., 2021)
• Foraging behaviour: cooperative (Greer et al., 2021)

Morphology

Worker Morphology

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• Antennal segment count: 11 • Antennal club: 3 • Palp formula: 5,3 • Total dental count: 5-6 (6 usual) • Spur formula: 0,0 • Eyes: 11->100 ommatidia • Pronotal Spines: absent • Mesonotal Spines: absent • Propodeal Spines: dentiform; present • Petiolar Spines: absent • Caste: none or weak • Sting: present • Metaplural Gland: present • Cocoon: absent

Male Morphology

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 • Antennal segment count 12 • Antennal club 0 • Palp formula 5,3 • Total dental count 0 • Spur formula 0, 0

Karyotype

Species Uncertain

• Leptothorax sp.1: n = 15 (Canada) (Loiselle et al., 1990).
• Leptothorax sp.1: n = 16 (Canada) (Loiselle et al., 1990).
• Leptothorax sp.1: n = 17 (USA) (Fischer, 1987).
• Leptothorax sp.2: n = 12 (Algeria) (Fischer, 1987).
• Leptothorax sp.3: n = 12 (Italy) (Fischer, 1987).
• Leptothorax sp.4: n = 13 (Croatia) (Fischer, 1987).
• Leptothorax sp.5: n = 14 (Canada) (Fischer, 1987).
• Leptothorax sp.6: n = 22 (France) (Fischer, 1987).
• Leptothorax sp.A: n = 17 (USA) (Loiselle et al., 1990).
• Leptothorax: n = 21 (France) (Fischer, 1987).
• Leptothorax: n = 16 (Japan) (Imai, 1969).

All Karyotype Records for Genus

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Taxon	Haploid	Diploid	Karyotype	Locality	Source	Notes
Leptothorax	12			Algeria	Fischer, 1987
Leptothorax	12			Italy	Fischer, 1987
Leptothorax	13			Croatia	Fischer, 1987
Leptothorax	14			Canada	Fischer, 1987
Leptothorax	15			Canada	Loiselle et al., 1990
Leptothorax	16			Canada	Loiselle et al., 1990
Leptothorax	16			Japan	Imai, 1969
Leptothorax	17			USA	Fischer, 1987
Leptothorax	17			USA	Loiselle et al., 1990
Leptothorax	21			France	Fischer, 1987
Leptothorax	22			France	Fischer, 1987
Leptothorax acervorum	13	26		Canada; France; Germany; Sweden; Switzerland	Fischer, 1987; Hauschteck-Jungen & Jungen, 1983; Loiselle et al., 1990
Leptothorax canadensis	16			Canada; USA	Loiselle et al., 1990
Leptothorax canadensis	17	34		Canada; Germany; USA	Buschinger et al., 1980; Fischer, 1987; Loiselle et al., 1990	possible variation due B chromosomes
Leptothorax canadensis	18	36		Canada; Germany; USA	Fischer, 1987; Loiselle et al., 1990	possible variation due B chromosomes
Leptothorax canadensis	22	44		Canada; Germany; USA	Fischer, 1987	possible variation due B chromosomes
Leptothorax canadensis	23			Canada; USA	Loiselle et al., 1990	possible variation due B chromosomes
Leptothorax crassipilis	17			USA	Francoeur, 1986; Loiselle et al., 1990
Leptothorax crassipilis	18			USA	Francoeur, 1986; Loiselle et al., 1990
Leptothorax faberi	15			Canada	Buschinger, 1982
Leptothorax faberi	16			Canada	Fischer, 1987
Leptothorax goesswaldi	28			France; Switzerland	Fischer, 1987	as ''Doronomyrmex goesswaldi''
Leptothorax gredleri	11			Germany; Switzerland	Buschinger et al., 1980; Fischer, 1987; Loiselle et al., 1990
Leptothorax kutteri	23 • 24 • 25			France; Germany; Italy; Sweden; Switzerland	Buschinger & Fischer, 1991; Douwes & Buschinger, 1983; Fischer, 1987	as ''Doronomyrmex kutteri''
Leptothorax muscorum	17	34		Canada; Germany; USA	Buschinger et al., 1980; Fischer, 1987; Loiselle et al., 1990	possible variation due B chromosomes
Leptothorax muscorum	18	36		Canada; Germany; USA	Fischer, 1987; Loiselle et al., 1990	possible variation due B chromosomes
Leptothorax muscorum	22	44		Canada; Germany; USA	Fischer, 1987	possible variation due B chromosomes
Leptothorax pacis	26			France; Switzerland	Buschinger & Fischer, 1991; Fischer, 1987	as ''Doronomyrmex pacis''
Leptothorax pocahontas	18			Canada	Fischer, 1987; Buschinger & Heinze, 1993	as ''Doronomyrmex pocahontas''
Leptothorax retractus	17			Canada	Francoeur, 1986; Loiselle et al., 1990
Leptothorax retractus	18			Canada	Francoeur, 1986; Loiselle et al., 1990
Leptothorax sphagnicola	13			Canada	Francoeur, 1986; Fischer, 1987; Loiselle et al., 1990	as ''Leptothorax sphagnicolus''

Phylogeny

Myrmicinae

Myrmicini ⊞(2 genera) ⊟ Manica  (5 species, 2 fossil species) Myrmica  (210 species, 18 fossil species) Pogonomyrmecini ⊞(3 genera) ⊟ Patagonomyrmex  (3 species, 0 fossil species) Hylomyrma  (30 species, 0 fossil species) Pogonomyrmex  (95 species, 1 fossil species) Stenammini ⊞(7 genera) ⊟ Stenamma  (85 species, 1 fossil species) Novomessor  (3 species, 0 fossil species) Veromessor  (10 species, 0 fossil species) some Aphaenogaster  (218 species, 12 fossil species) some Aphaenogaster Messor  (169 species, 1 fossil species) Goniomma  (10 species, 1 fossil species) Oxyopomyrmex  (12 species, 0 fossil species) Solenopsidini ⊞(22 genera) ⊟ Stegomyrmex  (5 species, 0 fossil species) Dolopomyrmex  (1 species, 0 fossil species) some Rogeria  (40 species, 0 fossil species) Bariamyrma  (1 species, 0 fossil species) some Rogeria Solenopsis  (215 species, 9 fossil species) Kempfidris  (1 species, 0 fossil species) Tropidomyrmex  (1 species, 0 fossil species) Austromorium  (2 species, 0 fossil species) some Monomorium  (320 species, 3 fossil species) some Monomorium Oxyepoecus  (21 species, 0 fossil species) Megalomyrmex  (45 species, 0 fossil species) Tyrannomyrmex  (4 species, 0 fossil species) some Monomorium Epelysidris  (1 species, 0 fossil species) Anillomyrma  (2 species, 0 fossil species) Myrmicaria  (71 species, 0 fossil species) some Monomorium Syllophopsis  (23 species, 0 fossil species) some Adelomyrmex  (31 species, 0 fossil species) Cryptomyrmex  (2 species, 0 fossil species) some Adelomyrmex Baracidris  (3 species, 0 fossil species) Attini ⊞(49 genera) ⊟ Ochetomyrmex  (2 species, 0 fossil species) Tranopelta  (2 species, 0 fossil species) Diaphoromyrma  (1 species, 0 fossil species) Lachnomyrmex  (16 species, 0 fossil species) Blepharidatta  (4 species, 0 fossil species) Allomerus  (8 species, 0 fossil species) Wasmannia  (11 species, 0 fossil species) Pheidole  (1,294 species, 7 fossil species) Cephalotes  (123 species, 16 fossil species) Procryptocerus  (44 species, 0 fossil species) Strumigenys  (879 species, 4 fossil species) Phalacromyrmex  (1 species, 0 fossil species) Pilotrochus  (1 species, 0 fossil species) Protalaridris  (7 species, 0 fossil species) Rhopalothrix  (19 species, 0 fossil species) Basiceros  (9 species, 0 fossil species) Octostruma  (35 species, 0 fossil species) Eurhopalothrix  (54 species, 0 fossil species) Talaridris  (1 species, 0 fossil species) Acanthognathus  (7 species, 1 fossil species) Daceton  (2 species, 0 fossil species) Lenomyrmex  (7 species, 0 fossil species) Microdaceton  (4 species, 0 fossil species) Orectognathus  (29 species, 0 fossil species) Colobostruma  (16 species, 0 fossil species) Epopostruma  (20 species, 0 fossil species) Mesostruma  (9 species, 0 fossil species) Paleoattina Apterostigma  (44 species, 2 fossil species) Mycocepurus  (6 species, 0 fossil species) Myrmicocrypta  (31 species, 0 fossil species) Neoattina Cyatta  (1 species, 0 fossil species) Kalathomyrmex  (1 species, 0 fossil species) Mycetarotes  (4 species, 0 fossil species) Mycetosoritis  (2 species, 0 fossil species) some Cyphomyrmex  (23 species, 2 fossil species) some Cyphomyrmex Paramycetophylax  (1 species, 0 fossil species) Mycetophylax  (21 species, 0 fossil species) Mycetagroicus  (4 species, 0 fossil species) Mycetomoellerius  (31 species, 1 fossil species) Sericomyrmex  (11 species, 0 fossil species) Xerolitor  (1 species, 0 fossil species) Paratrachymyrmex  (9 species, 0 fossil species) Trachymyrmex  (9 species, 0 fossil species) Amoimyrmex  (3 species, 0 fossil species) Atta  (20 species, 1 fossil species) some Acromyrmex  (56 species, 0 fossil species) some Acromyrmex Pseudoatta  (2 species, 0 fossil species) Crematogastrini ⊞(65 genera) ⊟ Rostromyrmex  (1 species, 6 fossil species) Cardiocondyla  (90 species, 0 fossil species) Ocymyrmex  (34 species, 0 fossil species) Nesomyrmex  (84 species, 2 fossil species) Xenomyrmex  (5 species, 0 fossil species) Terataner  (14 species, 0 fossil species) Atopomyrmex  (3 species, 0 fossil species) Cataulacus  (65 species, 3 fossil species) Carebara  (249 species, 9 fossil species) Diplomorium  (1 species, 0 fossil species) Melissotarsus  (4 species, 1 fossil species) Rhopalomastix  (14 species, 0 fossil species) Calyptomyrmex  (38 species, 0 fossil species) Strongylognathus  (27 species, 0 fossil species), Tetramorium  (598 species, 2 fossil species) Cyphoidris  (4 species, 0 fossil species) Dicroaspis  (2 species, 0 fossil species) Aretidris  (2 species, 0 fossil species) Vollenhovia  (83 species, 3 fossil species) Dacetinops  (7 species, 0 fossil species) Indomyrma  (2 species, 0 fossil species) Crematogaster  (783 species, 3 fossil species) Meranoplus  (91 species, 0 fossil species) Lophomyrmex  (13 species, 0 fossil species) Adlerzia  (1 species, 0 fossil species) Recurvidris  (12 species, 0 fossil species) Stereomyrmex  (3 species, 0 fossil species) Trichomyrmex  (29 species, 0 fossil species) Eutetramorium  (3 species, 0 fossil species) Royidris  (15 species, 0 fossil species) Malagidris  (6 species, 0 fossil species) Vitsika  (16 species, 0 fossil species) Huberia  (2 species, 0 fossil species) Podomyrma  (62 species, 1 fossil species) Liomyrmex  (1 species, 0 fossil species) Metapone  (31 species, 0 fossil species) Kartidris  (6 species, 0 fossil species) Mayriella  (9 species, 0 fossil species) Tetheamyrma  (2 species, 0 fossil species) Dacatria  (1 species, 0 fossil species) Proatta  (1 species, 0 fossil species) Dilobocondyla  (22 species, 0 fossil species) Secostruma  (1 species, 0 fossil species) Acanthomyrmex  (19 species, 0 fossil species) Myrmecina  (106 species, 0 fossil species) Perissomyrmex  (6 species, 0 fossil species) Pristomyrmex  (61 species, 3 fossil species) some Lordomyrma  (36 species, 0 fossil species) Propodilobus  (1 species, 0 fossil species) Lasiomyrma  (4 species, 0 fossil species) some Lordomyrma Ancyridris  (2 species, 0 fossil species) some Lordomyrma Paratopula  (12 species, 0 fossil species) Poecilomyrma  (2 species, 0 fossil species) Romblonella  (10 species, 0 fossil species) Rotastruma  (3 species, 0 fossil species) Gauromyrmex  (3 species, 0 fossil species) Vombisidris  (19 species, 0 fossil species) Temnothorax  (504 species, 7 fossil species) Harpagoxenus  (4 species, 0 fossil species) Formicoxenus  (8 species, 0 fossil species) Leptothorax  (20 species, 0 fossil species)

See Phylogeny of Myrmicinae for details.

Nomenclature

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

• LEPTOTHORAX [Myrmicinae: Formicoxenini]
  • Leptothorax Mayr, 1855: 431. Type-species: Formica acervorum, by subsequent designation of Bingham, 1903: 214.
  • [Type-species not Myrmica clypeata, unjustified subsequent designation by Emery, 1912d: 271; repeated in Wheeler, W.M. 1913a: 79 and Emery, 1924d: 248.]
  • Leptothorax senior synonym of Mychothorax: Smith, M.R. 1950: 29 [both have the same type-species, synonymy is therefore absolute].
  • Leptothorax senior synonym of Doronomyrmex: Heinze, 1998: 195.
• DORONOMYRMEX [junior synonym of Leptothorax]
  • Doronomyrmex Kutter, 1945: 485. Type-species: Doronomyrmex pacis, by monotypy.
  • Doronomyrmex junior synonym of Leptothorax: Heinze, 1998: 195.
• MYCHOTHORAX [junior synonym of Leptothorax]
  • Mychothorax Ruzsky, 1904a: 288. Type-species: Formica acervorum, by original designation.
  • Mychothorax subgenus of Leptothorax: Ruzsky, 1905b: 609; Emery, 1915g: 24.
  • Mychothorax junior synonym of Leptothorax: Smith, M.R. 1950: 29.
  • [Leptothorax and Mychothorax share the same type-species, synonymy is therefore absolute.]

References

• Arnold, G. 1916. A monograph of the Formicidae of South Africa. Part II. Ponerinae, Dorylinae. Ann. S. Afr. Mus. 14: 159-270 (page 257, Leptothorax in Myrmicinae, Leptothoracini)
• Ashmead, W. H. 1905c. A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. Can. Entomol. 37: 381-384 (page 383, Leptothorax in Myrmicinae, Stenammini)
• Barden, P. 2017. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages. Myrmecological News 24: 1-30.
• Bingham, C. T. 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. London: Taylor and Francis, 506 pp. (page 214, Type-species: Formica acervorum, by subsequent designatio)
• Blaimer, B.B., Ward, P.S., Schultz, T.R., Fisher, B.L., Brady, S.G. 2018. Paleotropical diversification dominates the evolution of the hyperdiverse ant tribe Crematogastrini (Hymenoptera: Formicidae). Insect Systematics and Diversity 2(5): 3; 1-14 (doi:10.1093/isd/ixy013).
• Bolton, B. 1982. Afrotropical species of the myrmecine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor and Cataulacus (Formicidae). Bulletin of the British Museum (Natural History). Entomology, 46: 307-370 (page 319, Leptothorax senior synonym of Dichothorax, Mychothorax, Myrafant (and its junior synonym Icothorax), Myrmammophilus, Nesomyrmex (and its junior synonyms Caulomyrma, Goniothorax (junior homonym) and Limnomyrmex), Temnothorqax, Tetramyrma)
• Bolton, B. 1994. Identification guide to the ant genera of the world. Cambridge, Mass.: Harvard University Press, 222 pp. (page 105, Leptothorax in Myrmicinae, Formicoxenini)
• Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 247, Leptothorax in Myrmicinae, Formicoxenini)
• Brown, W. L., Jr. 1973b. A comparison of the Hylean and Congo-West African rain forest ant faunas. Pp. 161-185 in: Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Washington, D.C.: Smithsonian Institution Press, viii + 350 pp. (page 178-185, Leptothorax provisional senior synonym of Antillaemyrmex, Caulomyrma, Croesomyrmex, Dichohorax, Goniothorax, Icothorax, Limnomyrmex, Macromischa, Mychothorax, Myrmammophilus, Myrafant, Nesomyrmex, Temnothorax)
• Burchill, A.T., Moreau, C.S. 2016. Colony size evolution in ants: macroevolutionary trends. Insectes Sociaux 63, 291–298 (doi:10.1007/s00040-016-0465-3).
• Cantone S. 2018. Winged Ants, The queen. Dichotomous key to genera of winged female ants in the World. The Wings of Ants: morphological and systematic relationships (self-published).
• Dalla Torre, K. W. von. 1893. Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann, 289 pp. (page 122, Leptothorax in Myrmicinae)
• Emery, C. 1877b. Saggio di un ordinamento naturale dei Mirmicidei, e considerazioni sulla filogenesi delle formiche. Bull. Soc. Entomol. Ital. 9: 67-83 (page 81, Leptothorax in Myrmicidae, Myrmicidae`)
• Emery, C. 1895l. Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 8: 685-778 (page 769, Leptothorax in Myrmicinae, Myrmicini)
• Emery, C. 1914e. Intorno alla classificazione dei Myrmicinae. Rend. Sess. R. Accad. Sci. Ist. Bologna Cl. Sci. Fis. (n.s.) 18: 29-42 (page 42, Leptothorax in Myrmicinae, Leptothoracini)
• Emery, C. 1922c. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [part]. Genera Insectorum 174B: 95-206 (page 248, Type-species not Myrmica clypeata, unjustified subsequent designation.)
• Emery, C. 1924f [1922]. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [concl.]. Genera Insectorum 174C: 207-397 (page 247, Leptothorax in Myrmicinae, Leptothoracini)
• Emery, C.; Forel, A. 1879. Catalogue des Formicides d'Europe. Mitt. Schweiz. Entomol. Ges. 5: 441-481 (page 458, Leptothorax in Myrmicinae [Myrmicidae])
• Forel, A. 1895b. A fauna das formigas do Brazil. Bol. Mus. Para. Hist. Nat. Ethnogr. 1: 89-139 (page 125, Leptothorax in Myrmicinae, Myrmicini)
• Forel, A. 1899d. Formicidae. [part]. Biol. Cent.-Am. Hym. 3: 25-56 (page 54, Leptothorax in Myrmicinae, Myrmicini)
• Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 244, Leptothorax in Myrmicinae, Leptothoracini)
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