Camponotus

Camponotus Temporal range: 53–0 Ma PreЄ Є O S D C P T J K Pg N Ypresian, Early Eocene – Recent
Camponotus ligniperda
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Formicinae
Tribe:	Camponotini
Genus:	Camponotus Mayr, 1861
Type species
Formica ligniperda, now Camponotus ligniperda
Subgenera
Camponotus Mayr, 1861 Dendromyrmex Emery, 1895 Forelophilus Kutter, 1931 Hypercolobopsis Emery, 1920 Karavaievia Emery, 1925 Manniella Wheeler, W.M., 1921 Mayria Forel, 1878 Myrmacrhaphe Santschi, 1926 Myrmamblys Forel, 1912 Myrmaphaenus Emery, 1920 Myrmentoma Forel, 1912 Myrmepinotus Santschi, 1921 Myrmepomis Forel, 1912 Myrmespera Santschi, 1926 Myrmeurynota Forel, 1912 Myrmisolepis Santschi, 1921 Myrmobrachys Forel, 1912 Myrmocladoecus Wheeler, W.M., 1921 Myrmodirachis Emery, 1925 Myrmogonia Forel, 1912 Myrmomalis Forel, 1914 Myrmonesites Emery, 1920 Myrmopalpella Stärcke, 1934 Myrmopelta Santschi, 1921 Myrmophyma Forel, 1912 Myrmopiromis Wheeler, W.M., 1921 Myrmoplatypus Santschi, 1921 Myrmoplatys Forel, 1916 Myrmopsamma Forel, 1914 Myrmopytia Emery, 1920 Myrmosaulus Wheeler, W.M., 1921 Myrmosericus Forel, 1912 Myrmosphincta Forel, 1912 Myrmostenus Emery, 1920 Myrmotarsus Forel, 1912 Myrmotemnus Emery, 1920 Myrmothrix Forel, 1912 Myrmotrema Forel, 1912 Myrmoxygenys Emery, 1925 Orthonotomyrmex Ashmead, 1906 Paramyrmamblys Santschi, 1926 Phasmomyrmex Stitz, 1910 Pseudocolobopsis Emery, 1920 Rhinomyrmex Forel, 1886 Tanaemyrmex Ashmead, 1905 Thlipsepinotus Santschi, 1928
Diversity
1,497 species 32 fossil species (Species Checklist, Species by Country) Camponotus ligniperda Specimen Label
Synonyms
Condylomyrma Santschi, 1928 Myrmocamelus Forel, 1914 Myrmolophus Emery, 1920 Myrmosaga Forel, 1912 Myrmoturba Forel, 1912 Neocolobopsis Borgmeier, 1928 Neomyrmamblys Wheeler, W.M., 1921 Orthonotus Ashmead, 1905 Paleosminthurus Pierce & Gibron, 1962 Paracolobopsis Emery, 1920 Shanwangella Zhang, J., 1989

Evolutionary Relationships

Gigantiops   (1 species) Santschiella   (1 species) Myrmoteras   (41 species) Camponotini Colobopsis   (117 species) Opisthopsis   (20 species) Dinomyrmex   (2 species) Polyrhachis   (782 species) Camponotus   (1,497 species) Calomyrmex   (14 species) Echinopla   (39 species)

Based on Ward et al. 2016.

Hita Garcia, Wiesel and Fischer (2013) - Camponotus is an extremely large and complex, globally distributed genus. At present, more than 1000 species and nearly 500 subspecies belonging to 45 subgenera are described (Bolton, 2012) and it could well be the largest ant genus of all. The enormous species richness, high levels of intraspecific and geographic variation and polymorphism render the taxonomy of Camponotus one of the most complex and difficult. Revisionary studies on Camponotus are generally confined to species groups and/or small geographical regions (e.g. Robertson & Zachariades, 1997; Snelling, 2006; McArthur, 2007; Shattuck & Janda, 2009). These ants live in a variety of habitats and microhabitats and the sheer size of the genus makes any characterisation of their biology challenging. Nests are built in the ground, in rotten branches or twigs, or rarely into living wood (Bolton, 1973a) and most species possess a highly generalistic diet.

Recently, Blaimer et al. (2015) found that Camponotus is paraphyletic with respect to Phasmomyrmex and Forelophilus, and that Colobopsis is sister to the remainder of the Camponotini while Dinomyrmex is nested outside of Camponotus. For this reason, Ward et al. (2016) revised the generic delimitation of the Camponotini (see the individual generic pages for additional information).

Photo Gallery

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  Camponotus pennsylvanicus feeding on a droplet of honey.
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  A queen from Dominican amber.
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  Mermithid nematode exposed in the gaster of Camponotus sp. from the Sierra Nevada, California (Poinar, 2012, Fig. 2).

Identification

Camponotus is a hyperdiverse genus (> 1,000 species, > 400 subspecies) that is rife with taxonomic problems and puzzles. Various subgenera, complexes, and informal groupings have either been defined in taxonomic publications or have been informally recognized. All of this sorting of species at the level between the genus and species level is an attempt to introduce some order to the genus and/or allow for a more workable framework for understanding Camponotus diversity within a particular region. The Camponotus species groups page provides links to some of the Antwiki pages that include Camponotus species groupings and information. See, particularly, Emery's (1925) keys to the subgenera of Camponotus, as well as related pages linked therein.

Note that female "Camponotus" are usually distinguishable from "Colobopsis" by their antennal sockets, which are placed in the anterior half of the frontal carinae (see the Colobopsis taxon account for more detail).

See images of species within this genus

Keys including this Genus

• Key to Ant Genera of the Navajo Reservation
• Key to Australian Genera of Formicinae
• Key to North American Genera of Formicinae
• Key to the Ant Genera of New Mexico

Keys to Subgenera or Species Groups in this Genus

• Genera Insectorum: Emery's key to Camponotus subgenera of the New World
• Genera Insectorum: Emery's key to Camponotus subgenera of the Old World

Keys to Species in this Genus

• Key to Australian Camponotus species
• Key to Camponotus aureopilis species-group
• Key to Camponotus of Turkey
• Key to Afrotropical Camponotus fulvopilosus species group
• Key to Camponotus of Israel
• Key to Camponotus Karavaievia workers
• Key to Camponotus Karavaievia queens
• Key to Camponotus Karavaievia males
• Key to Camponotus maculatus species complex in the New World
• Key to Australian Camponotus majors of the southwestern Botanical Province
• Key to Australian Camponotus minors of the southwestern Botanical Province
• Key to Malagasy Camponotus edmondi species group
• Key to Malagasy Camponotus grandidieri and niveosetosus species groups
• Key to Camponotus Myrmopytia workers
• Key to Malagasy Camponotus Mayria
• Key to Camponotus Myrmostenus Queens
• Key to the Westpalaearctic Camponotus lateralis species group

Emery's keys to the subgenera from Genera Insectorum

Emery's (1925) key to subgenera from Genera Insectorum

Species Groups

• Table of Camponotus species by subgenus
• Afrotropical species by subgenus, No images
• Afrotropical species by subgenus, with images
• Afrotropical species in Alphabetic order, with images
• Malagasy Camponotus edmondi species group
• Camponotus grandidieri species group
• Camponotus niveosetosus species group

Biology

A twig-nesting species from Cairns, Queensland, Australia.

Association with Other Organisms

Species Uncertain

Hymenoptera

Unspecified species within this genus are hosts for the following parasitic wasps (Universal Chalcidoidea Database):

Wasp Family	Wasp Species	Relationship
Encyrtidae	Arketypon vaderi	associate
Eucharitidae	Ancylotropus manipurensis	primary host
Eucharitidae	Lophyrocera variabilis	primary host
Eucharitidae	Mateucharis rugulosa	primary host
Eucharitidae	Pseudochalcura gibbosa	primary host
Eucharitidae	Pseudochalcura nigrocyanea	associate
Eucharitidae	Rhipipalloidea sp.	primary host
Eucharitidae	Rhipipalloidea madangensis	primary host
Eucharitidae	Stilbula cyniformis	primary host
Eucharitidae	Stilbuloida doddi	primary host
Eucharitidae	Zulucharis campbelli	primary host
Eulophidae	Horismenus microdonophagus	associate
Eulophidae	Horismenus myrmecophagus	primary host
Eurytomidae	Camponotophilus delvarei	associate

Fungi

Unknown species of Camponotus are hosts for the fungi Ophiocordyceps albacongiuae, Ophiocordyceps blakebarnesii, Ophiocordyceps rami and Ophiocordyceps septa (Araujo et al., 2018).

All Associate Records for Genus

Explore Associate Data: All, Drilldown

Taxon	Relationship	Associate Type	Associate Taxon	Associate Relationship	Locality	Source	Notes
Camponotus	host	encyrtid wasp	Arketypon vaderi	parasite		Universal Chalcidoidea Database	associate
Camponotus	host	eucharitid wasp	Ancylotropus manipurensis	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Lophyrocera variabilis	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Mateucharis rugulosa	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Pseudochalcura gibbosa	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Pseudochalcura nigrocyanea	parasite		Universal Chalcidoidea Database	associate
Camponotus	host	eucharitid wasp	Rhipipalloidea madangensis	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Rhipipalloidea sp.	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Stilbula cyniformis	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Stilbuloida doddi	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eucharitid wasp	Zulucharis campbelli	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eulophid wasp	Horismenus microdonophagus	parasite		Universal Chalcidoidea Database	associate
Camponotus	host	eulophid wasp	Horismenus myrmecophagus	parasite		Universal Chalcidoidea Database	primary host
Camponotus	host	eurytomid wasp	Camponotophilus delvarei	parasite		Universal Chalcidoidea Database	associate
Camponotus	host	fungus	Ophiocordyceps albacongiuae	pathogen		Araujo et al., 2018
Camponotus	host	fungus	Ophiocordyceps blakebarnesii	pathogen		Araujo et al., 2018
Camponotus	host	fungus	Ophiocordyceps rami	pathogen		Araujo et al., 2018
Camponotus	host	fungus	Ophiocordyceps septa	pathogen		Araujo et al., 2018
Camponotus aberrans	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus acvapimensis	host	encyrtid wasp	Anagyrus lopezi	parasite		Universal Chalcidoidea Database	associate
Camponotus aethiops	host	ant	Camponotus ruseni	inquiline		Karaman, 2012
Camponotus aethiops	host	ant	Camponotus universitatis	inquiline		Tinaut et al., 1992; Guillem et al., 2014
Camponotus aethiops	host	eucharitid wasp	Stilbula cyniformis	parasite		Universal Chalcidoidea Database	primary host
Camponotus aethiops	host	fungus	Laboulbenia camponoti	pathogen		Espadaler & Santamaria, 2012
Camponotus aethiops	host	nematode	Camponotimermis bifidus	parasite		Ipateva et al., 1990
Camponotus aethiops	host	nematode	Comanimermis clujensis	parasite		Coman, 1953
Camponotus aethiops	mutualist	butterfly	Glaucopsyche alexis			Obregon et al. 2015
Camponotus americanus	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus atriceps	host	''Microdon'' fly	Microdon fulgens	predator	Florida, Georgia (type), New Jersey, Pennsylvania, Texas	Wiedemann, 1830
Camponotus atriceps	host	eucharitid wasp	Obeza floridana	parasite		Universal Chalcidoidea Database	primary host
Camponotus atriceps	host	eulophid wasp	Horismenus floridensis	parasite		Universal Chalcidoidea Database	primary host
Camponotus atriceps	host	fungus	Ophiocordyceps camponoti-atricipis	pathogen		Araujo et al., 2018; Shrestha et al., 2017
Camponotus atriceps	host	fungus	Ophiocordyceps camponoti-atricipis	pathogen		Sobczak et al., 2017
Camponotus atriceps	host	fungus	Ophiocordyceps lloydii	pathogen		Araujo et al., 2018; Shrestha et al., 2017
Camponotus atriceps	host	fungus	Ophiocordyceps unilaterialis	pathogen		Araujo et al., 2018; Shrestha et al., 2017
Camponotus baldaccii	host	fungus	Laboulbenia camponoti	pathogen		Espadaler & Santamaria, 2012
Camponotus balzani	host	fungus	Ophiocordyceps camponoti-balzani	pathogen		Araujo et al., 2018; Evans et al., 2011; Shrestha et al., 2017
Camponotus bispinosus	host	fungus	Ophiocordyceps camponoti-bispinosi	pathogen		Araujo et al., 2018; Shrestha et al., 2017
Camponotus brutus	host	encyrtid wasp	Ooencyrtus sp.	parasite		Universal Chalcidoidea Database	associate
Camponotus brutus	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus buchneri	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus castaneus	host	fungus	Ophiocordyceps kimflemingiae	pathogen		Araujo et al., 2018
Camponotus castaneus	host	nematode	Rabbium paradoxus	parasite	Florida	Poinar et al., 1989
Camponotus chartifex	host	fungus	Ophiocordyceps camponoti-chartificis	pathogen		Araujo et al., 2018
Camponotus compressus	host	chalcid wasp	Epitranus chilkaensis	parasite		Universal Chalcidoidea Database	primary host
Camponotus compressus	mutualist	butterfly	Lampides boeticus			Obregon et al. 2015
Camponotus cruentatus	host	fungus	Ophiocordyceps halabalaensis	pathogen		Shrestha et al., 2017
Camponotus cruentatus	host	fungus	Ophiocordyceps ridleyi	pathogen		Shrestha et al., 2017
Camponotus cruentatus	mutualist	butterfly	Glaucopsyche alexis			Obregon et al. 2015
Camponotus cruentatus	mutualist	butterfly	Lampides boeticus			Obregon et al. 2015
Camponotus fallax	mutualist	butterfly	Polyommatus celina			Obregon et al. 2015
Camponotus femoratus	host	fungus	Ophiocordyceps camponoti-femorati	pathogen		Araujo et al., 2018
Camponotus flavomarginatus	host	encyrtid wasp	Anagyrus lopezi	parasite		Universal Chalcidoidea Database	associate
Camponotus floridanus	host	fungus	Ophiocordyceps camponoti-floridani	pathogen		Araujo et al., 2018
Camponotus foreli	mutualist	butterfly	Lampides boeticus			Obregon et al. 2015
Camponotus herculeanus	host	''Microdon'' fly	Microdon piperi	predator	Oregon, Montana, Washington (type)	Knab, 1917
Camponotus herculeanus	host	eucharitid wasp	Stilbula cyniformis	parasite		Universal Chalcidoidea Database	primary host
Camponotus herculeanus	host	nematode	Oscheius dolichurus	parasite	Ontario	Nickle & Ayre, 1966
Camponotus hippocrepis	host	fungus	Ophiocordyceps camponoti-hippocrepidis	pathogen		Araujo et al., 2018
Camponotus japonicus	host	ant	Polyrhachis lamellidens	temporary parasite		Kohriba, 1963
Camponotus japonicus	host	butterfly	Lampides argyrognomon	myrmecophile		Mizuno et al., 2018
Camponotus japonicus	host	fungus	Ophiocordyceps formicarum	pathogen		Shrestha et al., 2017
Camponotus kiusiuensis	host	ant	Polyrhachis lamellidens	temporary parasite		Sakai, 1990	needs confirmation
Camponotus laevigatus	host	''Microdon'' fly	Microdon piperi	predator	Oregon, Montana, Washington (type)	Knab, 1917
Camponotus ligniperda	host	fungus	Desmidiospora myrmecophila	pathogen		based on a photo by Michal Kukla, fungal identification by João Araújo
Camponotus ligniperda	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus maculatus	host	eucharitid wasp	Stilbula cyniformis	parasite		Universal Chalcidoidea Database	primary host
Camponotus melanoticus	host	fungus	Ophiocordyceps camponoti-melanotici	pathogen		Araujo et al., 2018; Evans et al., 2011; Shrestha et al., 2017
Camponotus modoc	host	''Microdon'' fly	Microdon albicomatus	predator	Idaho (type), Washington	Novak, 1977
Camponotus modoc	host	''Microdon'' fly	Microdon piperi	predator	Oregon, Montana, Washington (type)	Knab, 1917
Camponotus nidulans	host	fungus	Ophiocordyceps camponoti-nidulantis	pathogen		Araujo et al., 2018
Camponotus novaeboracensis	host	eucharitid wasp	Pseudochalcura gibbosa	parasite		Universal Chalcidoidea Database	primary host
Camponotus novogranadensis	host	fungus	Ophiocordyceps camponoti-novogranadensis	pathogen		Araujo et al., 2018; Evans et al., 2011; Shrestha et al., 2017
Camponotus nylanderi	host	aphelinid wasp	Aphytis melinus	parasite		Universal Chalcidoidea Database	associate
Camponotus nylanderi	host	encyrtid wasp	Leptomastix dactylopii	parasite		Universal Chalcidoidea Database	associate
Camponotus obscuripes	host	ant	Polyrhachis lamellidens	temporary parasite		Sakai, 1990
Camponotus obscuripes	host	fungus	Ophiocordyceps formicarum	pathogen		Shrestha et al., 2017
Camponotus obscuripes	host	fungus	Ophiocordyceps pulvinata	pathogen		Shrestha et al., 2017
Camponotus obscuripes	host	fungus	Ophiocordyceps sessilis	pathogen		Shrestha et al., 2017
Camponotus paradoxus	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus pennsylvanicus	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus piceus	host	fungus	Ophiocordyceps myrmecophila	pathogen		Shrestha et al., 2017
Camponotus piceus	mutualist	butterfly	Glaucopsyche alexis			Obregon et al. 2015
Camponotus pilicornis	host	ant	Camponotus universitatis	inquiline		Tinaut et al., 1992; Guillem et al., 2014
Camponotus pilicornis	host	fungus	Laboulbenia camponoti	pathogen		Espadaler & Santamaria, 2012
Camponotus pilicornis	mutualist	butterfly	Glaucopsyche alexis			Obregon et al. 2015
Camponotus pilicornis	mutualist	butterfly	Tomares ballus			Obregon et al. 2015
Camponotus planatus	host	eucharitid wasp	Pseudochalcura sculpturata	parasite		Universal Chalcidoidea Database	primary host
Camponotus renggeri	host	fungus	Ophiocordyceps camponoti-renggeri	pathogen		Araujo et al., 2018
Camponotus rufipes	host	fungus	Ophiocordyceps camponoti-rufipedis	pathogen		Andersen et al., 2012; Araujo et al., 2018; Evans et al., 2011; Loreto et al., 2014; Shrestha et al., 2017
Camponotus ruseni	inquiline	ant	Camponotus aethiops			Karaman, 2012
Camponotus sanctaefidei leonhardi	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus sericeiventris	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017
Camponotus sexguttatus	host	fungus	Ophiocordyceps camponoti-sexguttati	pathogen		Araujo et al., 2018
Camponotus simillimus indianus	host	fungus	Ophiocordyceps camponoti-indiani	pathogen		Araujo et al., 2018; Shrestha et al., 2017
Camponotus sylvaticus	host	fungus	Laboulbenia camponoti	pathogen		Espadaler & Santamaria, 2012
Camponotus sylvaticus	mutualist	butterfly	Lampides boeticus			Obregon et al. 2015
Camponotus universitatis	host	fungus	Laboulbenia camponoti	pathogen		Espadaler & Santamaria, 2012
Camponotus universitatis	inquiline	ant	Camponotus aethiops			Tinaut et al., 1992
Camponotus universitatis	inquiline	ant	Camponotus pilicornis			Tinaut et al., 1992
Camponotus vicinus	host	nematode	Formicitylenchus oregonensis	parasite	Oregon, USA	Poinar, 2003
Camponotus vividus	host	fungus	Ophiocordyceps unilaterialis	pathogen		Shrestha et al., 2017

Morphology

Worker Morphology

 • Antennal segment count 12 • Antennal club absent-gradual • Palp formula 6,4; 5,4 • Total dental count 4-9 • Spur formula 1 simple, 1 simple; 1 simple-barbulate, 1 simple-pectinate; 1 simple-pectinate, 1 simple-pectinate; 0, 0 • Eyes present • Scrobes absent • Caste polymorphic • Sting absent

Karyotype

Species Uncertain

• Camponotus (Myrmobrachys) sp.: 2n = 40, karyotype = 4M+36A+13B (Brazil) (Mariano et al., 2001) (variation observed for 2n=41-43 due B-chromsomes).
• Camponotus sp.(ANIC-1): n = 23, 2n = 46 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-2): n = 23 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-3): 2n = 48 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-5): 2n = 32 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-8): 2n = 32 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-9): 2n = 38 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-10): 2n = 46 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-11): 2n = 32 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-12): 2n = 38 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-13): n = 10, 2n = 20 (Australia) (Imai et al., 1977).
• Camponotus sp.(ANIC-14): n = 19, 2n = 38 (Australia) (Imai et al., 1977).
• Camponotus sp.1: 2n = 39 (Indonesia) (Imai et al., 1985) (mentined as a result of Robertsonian polymorphism).
• Camponotus sp.1: n = 9, 2n = 18 (Japan) (Imai & Kubota, 1972) ('Myrmamblys group).
• Camponotus sp.1: 2n = 38 (Indonesia) (Imai et al., 1985).
• Camponotus sp.1: n = 19 (Malaysia) (Goni et al., 1982).
• Camponotus sp.1: 2n = 40 (Sarawak) (Tjan et al., 1986).
• Camponotus sp.2: 2n = 38 (Indonesia) (Imai et al., 1985).
• Camponotus sp.2: n = 20, 2n = 40 (Malaysia) (Goni et al., 1982; Imai et al., 1983).
• Camponotus sp.2: 2n = 20 (Sarawak) (Tjan et al., 1986).
• Camponotus sp.3: 2n = 38 (Malaysia) (Goni et al., 1982).
• Camponotus sp.4: n = 18, 2n = 36 (Malaysia) (Goni et al., 1982).
• Camponotus sp.5: n = 20, 2n = 40 (Malaysia) (Goni et al., 1982).
• Camponotus sp.6: 2n = 44 (Malaysia) (Goni et al., 1982).
• Camponotus sp.7: 2n = 32, karyotype = 10M+22A (India) (Imai et al., 1984) (near C. variegatus).
• Camponotus sp.7: n = 20 (Malaysia) (Imai et al., 1983).
• Camponotus sp.8: 2n = 38 (Malaysia) (Imai et al., 1983).
• Camponotus sp.8: 2n = 39 (Malaysia) (Imai et al., 1983).
• Camponotus sp.9: 2n = 34 (India) (Imai et al., 1984) (near C. variegatus).
• Camponotus sp.9: 2n = 35 (India) (Imai et al., 1984) (near C. variegatus).
• Camponotus sp.9: 2n = 40 (Malaysia) (Imai et al., 1983).
• Camponotus sp.10: 2n = 20, karyotype = 20M (India) (Imai et al., 1984) (near C. infuscus).
• Camponotus sp.10: n = 13 (Malaysia) (Imai et al., 1983).
• Camponotus sp.10: n = 14 (Malaysia) (Imai et al., 1983).
• Camponotus sp.11: 2n = 52 (Malaysia) (Imai et al., 1983).
• Camponotus sp.12: 2n = 34, karyotype = 10M+24A (India) (Imai et al., 1984) (near C. variegatus).
• Camponotus sp.12: 2n = 18 (Malaysia) (Imai et al., 1983).
• Camponotus sp.13: 2n = 18 (Malaysia) (Imai et al., 1983).
• Camponotus: n = 26, 2n = 52 (USA) (Crozier, 1970b) ('C. impressus group).
• Camponotus: 2n = 20 (Taiwan) (Hung et al., 1972) ('C. variegatus complex).
• Camponotus: n = 9, 2n = 18 (Japan) (Imai, 1966; Imai & Yosida, 1964).

All Karyotype Records for Genus

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Taxon	Haploid	Diploid	Karyotype	Locality	Source	Notes
Camponotus		18		Malaysia	Imai et al., 1983
Camponotus		20		Sarawak	Tjan et al., 1986
Camponotus		20		Taiwan	Hung et al., 1972	'C. variegatus'' complex
Camponotus		20	20M	India	Imai et al., 1984	near ''C. infuscus''
Camponotus		32		Australia	Imai et al., 1977
Camponotus		32	10M+22A	India	Imai et al., 1984	near ''C. variegatus''
Camponotus		34		India	Imai et al., 1984	near ''C. variegatus''
Camponotus		34	10M+24A	India	Imai et al., 1984	near ''C. variegatus''
Camponotus		35		India	Imai et al., 1984	near ''C. variegatus''
Camponotus		38		Australia	Imai et al., 1977
Camponotus		38		Indonesia	Imai et al., 1985
Camponotus		38		Malaysia	Goni et al., 1982
Camponotus		38		Malaysia	Imai et al., 1983
Camponotus		39		Indonesia	Imai et al., 1985	mentined as a result of Robertsonian polymorphism
Camponotus		39		Malaysia	Imai et al., 1983
Camponotus		40		Malaysia	Imai et al., 1983
Camponotus		40		Sarawak	Tjan et al., 1986
Camponotus		40	4M+36A+13B	Brazil	Mariano et al., 2001	variation observed for 2n=41-43 due B-chromsomes
Camponotus		44		Malaysia	Goni et al., 1982
Camponotus		46		Australia	Imai et al., 1977
Camponotus		48		Australia	Imai et al., 1977
Camponotus		52		Malaysia	Imai et al., 1983
Camponotus	10	20		Australia	Imai et al., 1977
Camponotus	13			Malaysia	Imai et al., 1983
Camponotus	14			Malaysia	Imai et al., 1983
Camponotus	18	36		Malaysia	Goni et al., 1982
Camponotus	19			Malaysia	Goni et al., 1982
Camponotus	19	38		Australia	Imai et al., 1977
Camponotus	20			Malaysia	Imai et al., 1983
Camponotus	20	40		Malaysia	Goni et al., 1982; Imai et al., 1983
Camponotus	20	40		Malaysia	Goni et al., 1982
Camponotus	23			Australia	Imai et al., 1977
Camponotus	23	46		Australia	Imai et al., 1977
Camponotus	26	52		USA	Crozier, 1970b	'C. impressus'' group
Camponotus	9	18		Japan	Imai & Kubota, 1972	'Myrmamblys'' group
Camponotus	9	18		Japan	Imai, 1966; Imai & Yosida, 1964
Camponotus aethiops		42		France; old Yugoslavia	Hauschteck-Jungen & Jungen, 1983
Camponotus alii		42		Spain	Hauschteck-Jungen & Jungen, 1983	Lorite and Palomeque 2010 states some issues about the identification/sampling location
Camponotus atriceps	20	40	4M+36A	Brazil	Mariano et al., 2001
Camponotus balzani	20	40	6M+34A	Brazil	Mariano et al., 2001
Camponotus barbatus taylori		24	24A	India	Imai et al., 1984	as ''Camponotus taylori''
Camponotus bonariensis		40	8M+32A	Uruguay	Goni et al., 1983
Camponotus cingulatus		40	4M+36A	Brazil	Mariano et al., 2001
Camponotus compressus	10 • 20	40		India; Tunisia	Hauschteck-Jungen & Jungen, 1983; Kumbkarni, 1965	Lorite and Palomeque 2010 states some issues about the identification/sampling location
Camponotus consobrinus	23	46		Australia	Imai et al., 1977
Camponotus crassisquamis		39 • 40	39A+1M; 40A	India	Imai et al., 1984	Lorite and Palomeque 2010 states the presence of heteromorphic chromosome in 2n=39
Camponotus crassus	10	20	20M	Brazil	Aguiar et al., 2016; Mariano et al., 2001
Camponotus cruentatus	18 • 20		12SM+5ST+3T	Spain	Hauschteck-Jungen & Jungen, 1983; Lorite et al., 2002b
Camponotus dolendus		20	20M	India	Imai et al., 1984
Camponotus femoratus		44	4ST+40T	French Guiana	Mariano et al., 2011
Camponotus festinus	19	38		Malaysia	Imai et al., 1983
Camponotus foreli	17	34		Spain	Hauschteck-Jungen & Jungen, 1983
Camponotus japonicus	13 • 14	26		Japan	Imai, 1966; Imai, 1969; Imai & Kubota, 1972; Imai & Yosida, 1964	In Imai et al. 1966 as ''Camponotus'' sp.; Lorite and Palomeque 2010 states that this karyotype could be a mistake
Camponotus kiusiuensis		28		Japan	Imai & Yosida, 1964; Imai, 1969; Imai, 1969
Camponotus lateralis		28		Switzerland	Hauschteck, 1962
Camponotus ligniperda	14	28		Switzerland	Hauschteck, 1961; Hauschteck-Jungen & Jungen, 1983
Camponotus mitis		20	20M	India	Imai et al., 1984
Camponotus mus		26	16SM+10A	Uruguay	Goni et al., 1983
Camponotus obscuripes		28		Japan	Imai, 1969
Camponotus parius		40	40A	India	Imai et al., 1984
Camponotus pilicornis		50		Spain	Hauschteck-Jungen & Jungen, 1983
Camponotus punctulatus		40	2SM+2ST+36A	Uruguay	Goni et al., 1983
Camponotus renggeri		40		Brazil	Aguiar et al., 2016
Camponotus rufipes	20	39 • 40	2SM+2ST+36A; 3M+36A; 4M+36A	Brazil; Uruguay	Aguiar et al., 2016; Goni et al., 1983; Mariano et al., 2001	Lorite and Palomeque 2010 states the presence of heteromorphic chromosome in 2n=39
Camponotus rufoglaucus	18			Spain; Tunisia	Hauschteck-Jungen & Jungen, 1983	Lorite and Palomeque 2010 states some issues about the identification/sampling location
Camponotus sericeiventris		40	4M+36A	Brazil	Mariano et al., 2001
Camponotus sericeus		44	2M+42A	India	Imai et al., 1984
Camponotus sylvaticus	20	40		Greece; Spain	Hauschteck-Jungen & Jungen, 1983
Camponotus thraso		40	40A	India	Imai et al., 1984
Camponotus vagus		28		Switzerland	Hauschteck, 1961
Camponotus variegatus		26	20M+6A	India	Imai et al., 1984
Camponotus vitiosus	9	18		Japan	Imai, 1966; Imai, 1969; Imai & Kubota, 1972	In Imai 1966a as ''Camponotus'' sp. and in Imai 1969 as ''C. caryae'' group sp. but the authors correct the identification by ''C. tokioensis'' and the current valid name is ''C. vitiosus''

Nomenclature

The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.

• CAMPONOTUS [Formicinae: Camponotini]
  • Camponotus Mayr, 1861: 35. Type-species: Formica ligniperda, by subsequent designation of Bingham, 1903: 347.
  • [Type-species not Formica herculeana, unjustified subsequent designation by Forel, 1914a: 259, repeated in Arnold, 1922: 612.]
  • Camponotus (subgenus not given) senior synonym of †Shanwangella: Hong & Wu, 2000: 20.
  • Camponotus senior synonym of †Paleosminthurus: Snelling, R.R. (pers. comm., unpublished).
  • Subgenera of Camponotus (alphabetical order with synonyms indicated): nominal plus Colobopsis (= Campylomyrma, = Condylomyrma, = Dolophra), Dendromyrmex, Dinomyrmex (= Myrmogigas), Hypercolobopsis (= Neocolobopsis), Karavaievia, Manniella, Mayria (= Myrmosaga), Myrmacrhaphe, Myrmamblys, Myrmaphaenus (= Neomyrmamblys, = Paracolobopsis), Myrmentoma, Myrmepinotus, Myrmepomis (= Myrmolophus), Myrmespera, Myrmeurynota, Myrmisolepis, Myrmobrachys, Myrmocladoecus, Myrmodirachis, Myrmogonia, Myrmomalis, Myrmonesites (= Myrmensites), Myrmopalpella, Myrmopelta, Myrmophyma (= Myrmocamelus), Myrmopiromis, Myrmoplatypus, Myrmoplatys, Myrmopsamma, Myrmopytia, Myrmosaulus, Myrmosericus, Myrmosphincta, Myrmostenus, Myrmotarsus, Myrmotemnus, Myrmothrix, Myrmotrema, Myrmoxygenys, Orthonotomyrmex (= Orthonotus (junior homonym), Paramyrmamblys, Pseudocolobopsis, Rhinomyrmex, Tanaemyrmex (= Myrmoturba), Thlipsepinotus.
  • [All subgenera were given as provisional junior synonyms of Camponotus by Brown, 1973b: 179-185. The list was repeated in Hölldobler & Wilson, 1990: 18 with all subgenera listed as junior synonyms. They reverted to subgeneric status in Bolton, 1994: 50; see under individual entries. The entry of Myrmophyma and Thlipsepinotus under the synonymy of Camponotus by Taylor & Brown, D.R. 1985: 109, is not accepted as confirmation as not all taxa were included.]
• †PALEOSMINTHURUS [junior synonym of Camponotus]
  • †Paleosminthurus Pierce & Gibron, 1962: 146. Type-species: †Paleosminthurus juliae (junior synonym of Formica festinata) [Snelling, R.R. pers. com., unpublished], by monotypy [in order Collembola].
  • †Paleosminthurus incertae sedis in Formicidae: Najt, 1987: 152; Bolton, 2003: 76.
  • †Paleosminthurus junior synonym of Camponotus: Snelling, R.R. (pers. comm., unpublished).
• †SHANWANGELLA [junior synonym of Camponotus]
  • †Shanwangella Zhang, J. 1989: 307. Type-species: †Shanwangella palaeoptera, by original designation.
  • †Shanwangella junior synonym of Camponotus: Hong & Wu, 2000: 20.

Common Names for Camponotus Subgenera

A mnemonic list of common names for the subgenera of Camponotus may be found here.

Subgenus Camponotus

True Carpenter Ants

Diagnosis of subgenus: Emery (1925) - "Worker and queen. - Large species; caste polymorphism and other characteristics as in the subgenus Tanaemyrmex, except for the following differences: Head of minors Alpha-form (see Tanaemyrmex for head shape classification), but head generally shorter and posterior head margin straighter; head of major worker much less enlarged and less emarginate posteriorly; head of queens similar to those of majors. Clypeus lacking medial carina nor usually with anterior lobe, although weakly-produced and more-or-less rectangular (C. japonicus, Mayr) or rounded (C. sansabeanus, Buckley) lobe may be present [note: C. sansabeanus is currently classified as Tanaemyrmex]; anterior margin of clypeus entire, lacking medial notch. Mandibles strongly arched, usually with 4 or 5 teeth, sometimes with 6. Mesosomal dorsum arched, always continuous in profile; pronotal dorsum rounded or sometimes depressed in majors, with slightly prominent humeri. Male. - Caracteres of Tanaemyrmex.

Note that the subgenera Camponotus, Tanaemyrmex, Myrmosericus, and Myrmothrix constitute a group: they are separated by characters of little importance and often offer transitions from one group to another.

Geographical distribution of species. - Holarctic region, except Mediterranean Africa; C. punctatissimus, Emery, is found in the north of Indomalaya."

(Translated and edited by B. E. Boudinot, 15 February 2017.)

Taxonomic History of Camponotus

The taxonomic history of Camponotus may be divided into four phases, or eras, based on the kind of work published during these time periods.

Phase I ("Archaic Era")

1861–1896 (Beginning: Mayr 1861; end: Emery 1896).

The beginning of subgeneric work on Camponotus started with Mayr's (1861) key to the European ant fauna, wherein he recognized the cork-head ants, Colobopsis, as a distinct genus for the first time. Most of the work during this period was regional in nature, treating Europe, the Mediterranean, Madagascar, Russia, India, Sri Lanka, and North America. Only one explicit subgenus, Mayria was designated, while two taxa currently treated as subgenera were described: the American weaving ants, Dendromyrmex, and big-nose carpenter ants, Rhinomyrmex. In 1896, Emery proposed the first global, synoptic classification of Camponotus; previously no higher classification was in place, excepting recognition Mayria. In this work, all of the groups Emery distinguished were treated informally. Specifically, Emery recognized three principal groups (cohorts), which he divided into 26 subgroups which he termed manipoli. This work marked the beginning of the second phase of higher classification within Camponotus, wherein formal treatments of these groups were published.

Phase II ("Classical Era")

1896–1928 (First classificatory publication of era for Camponotus: Emery, 1898; last: Santschi, 1928).

In addition to the continuation of global exploration marked by regional works (South America, India, Sri Lanka, Myanmar, Palearctic [numerous], North America, Afrotropics [particularly South Africa], and the Malagasy region), this period is distinguished by greater focus on the subgenera, with Forel (1914), Emery (1920), Wheeler (1922), Santschi (1921), and Arnold (1922) expanding upon and refining Emery's system. In 1912, Forel formalized the Emery's (1896) higher classification by recognizing 16 subgenera, but without designating types. Wheeler (1913) sought to correct this by explicitly designating type species, but unfortunately this work was overlooked when Forel (1914) redundantly designated type specimens for his subgenera. This error of omission was replicated by Emery (1920), wherein he revised his initial classification, recognizing a total of 34 subgenera, as well as excluding Phasmomyrmex and species belonging to Notostigma (Melophorini), which he described as new. Wheeler (1921) put the nail in the coffin for the issue of confusion over subgeneric type species in a pointedly titled article, Professor Emery's subgenera of the genus Camponotus Mayr. The capstone to this second phase was Emery' (1925) final ,monumental contribution to Wytsmann's Genera Insectorum, which is still the most significant work on the internal classification of Camponotus to date. In this work, Emery synthesized all previous taxonomic works on Camponotus, particularly those of Forel and Wheeler, revised the delimitation of all subgenera, including subgroups, and provided keys to and diagnoses of all subgenera, all of which amounted to an effort which has not been matched. Moreover, Emery proposed his final phylogenetic hypothesis for the genus (see Emery's Phylogenetic Hypothesis below), which is distinguished from hypotheses he proposed for other groups by the strong biogeographic structure he postulated for the group. In 1926 and 1928, Santschi published the final two works formally altering Emery's (1925) system; both of these works were minor acts of splitting, and neither revised Emery's keys.

Phase III ("Modern Era")

1928–2006 (Beginning: Santschi, 1928; end: Moreau et al. and Brady et al. 2006)

After Emery (1925) and Santschi (1926, 1928), no further works were published which formally modified the subgeneric system of Camponotus until the fourth era of the internal classification of the genus (see Phase IV below). For almost 90 years, there has been, effectively, no major attempt to understand the diversity of Camponotus globally, with the major publications of the era for the genus being regional catalogs (Kempf 1972 Neotropics (followed by Brandao’s update in 1991), Taylor & Brown 1985 Australia (followed by Shattuck 1999), and Bolton’s 1995 global catalog). Virtually all of the taxonomic work on Camponotus during this time were regional, which in sum span the globe, but with global revisions restricted to minor groups of species (i.e., the groups or complexes of herculeanus, nigriceps, fulvopilosus, weiderkehri, and perjerus), as well as regionally-restricted and well-defined or easily-recognizable subgenera (e.g., Dendromyrmex, Karavaievia, Myrmentoma, Myrmostenus, Myrmothrix). In the late ‘90s and early 2000’s, Shattuck and McArthur treated a number Australian groups which they recognized morphologically but in which they disregarded the subgeneric classification. The end of this era is marked by the first global phylogenies of the Formicidae based on molecular data, published by Moreau et al. (2006) and Brady et al. (2006).

Phase IV ("Molecular Era")

2006–present.

The current era of classification, with respect to Camponotus, is nascent. Blaimer et al. (2015) published two phylogenies of the Formicinae using “traditional” Sanger sequencing and “next-gen” Illumina sequencing of ultraconserved elements (UCEs). This work was followed shortly by a formal reclassification of the subfamily by Ward et al. (2016), in which a number of taxonomic changes were made to relieve Camponotus and particular subgenera of paraphyly (i.e., Colobopsis and Dinomyrmex revived status as genera, Myrmogonia synonymized with Colobopsis, Phasmomyrmex made subgenus of Camponotus).Bill MacKay is still working on his revision of Camponotus of the Americas, and at present, no major phylogenetic work is underway for Camponotus, beyond refining the placement of species belonging to Colobopsis.

(Section by B. E. Boudinot, 19 February 2017.)

Emery's Phylogenetic Hypothesis for Camponotus

In Emery's (1925) treatment of Camponotus in Genera Insectorum, he proposed a final, revised hypothesis of the internal relationships of the genus; an earlier version was published in Emery (1920). An important facet of this hypothesis, translated from p. 61, is Emery's "opinion that Camponotus underwent independent evolution in the Old and New Worlds". Emery postulated seven genus groups within Camponotus, with explicit morphologically-transitional lineages. These groups are indicated below:

Emery's original (1925) phylogenetic diagram.

Group I: Myrmopiromis

(Afrotropical and Malagasy regions, a few species from India)

1. Myrmopiromis

2. Myrmotrema

3. Myrmisolepis

4. Orthonotomyrmex

5. Myrmepinotus

Group II: Myrmosaulus

(Malaysia, Australia, Madagascar)

1. Myrmosaulus

Group III: Myrmamblys

Myrmopsamma subgroup (Africa)

1. Myrmopsamma

Karavaievia subgroup (Indomalaya, etc.)

1. Karavaievia

2. Myrmotarsus

3. Myrmoplatys

Mayria subgroup (Afrotropical, Malagasy)

1. Mayria

2. Myrmonesites

3. Myrmopytia

Miscellaneous subgroup

1. Myrmentoma

2. Colobopsis [note: revived status as genus, Ward et al. (2015)]

3. Myrmogonia [note: synonymized with Colobopsis, Ward et al. (2015)]

Subgroup transitional to Myrmaphaenus via Tanaemyrmex

1. Myrmamblys (Old World) [note: split into Myrmespera, Myrmopelta, Myrmotemnus, Paramyrmamblys, and Myrmamblys sensu stricto by Santschi (1926).]

2. Myrmophyma [note: split into Thlipsepinotus and Myrmophyma sensu stricto by Santschi (1928).

Group IV: Tanaemyrmex

(Cosmopolitan)

1. Tanaemyrmex [note: Emery's species groups may be particularly useful for understanding, and dividing, this probably "plesiotypic" taxon]

Group V: Myrmoxygenys group

(Transitional between Myrmamblys and Myrmaphaenus groups via Tanaemyrmex)

1. Myrmoxygenys (Afrotropics)

2. Dinomyrmex (Malaysia) [note: revived status as genus, Ward et al. (2015)]

3. Myrmosericus (Afrotropics, India)

4. Myrmothrix (Neotropics)

Group VI: Myrmaphaenus

Subgroup transitional to Myrmamblys group via Tanaemyrmex

1. Camponotus (Holarctic) (sensu lato, including Nearctic Tanaemyrmex and caryae and lateralis groups of Myrmentoma, in part.

2. Myrmaphaenus (New World)

Neotropical subgroup, transitional to Myrmobrachys group

1. Pseudocolobopsis

2. Myrmostenus

3. Hypercolobopsis

4. Manniella

5. Myrmosphincta

Group VII: Myrmobrachys

Subgroup transitional to Myrmaphaenus group

1. Myrmobrachys

2. Myrmocladoecus

3. Myrmomalis

4. Myrmeurynota

5. Myrmodirachis

6. Myrmoplatypus

7. Myrmepomis

(Section by B. E. Boudinot, 19 February 2017.)

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