Camponotus
Camponotus Temporal range: 53–0 Ma Ypresian, Early Eocene – Recent | |
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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 | |
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Diversity | |
1,497 species 30 fossil species (Species Checklist, Species by Country) | |
Synonyms | |
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Evolutionary Relationships | ||||||||||||||||||||||||||||||||||||||||||||||||||||||
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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).
Contents
Photo Gallery
- Camponotus pennsylvanicus feeding on a droplet of honey.
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).
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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
- Key to the New World Camponotus species complexes
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
- Key to Camponotus of Japan
- Key to Camponotus kiesenwetteri species group
- Key to Camponotus samius species complex
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
Distribution
Distribution and Richness based on AntMaps
Fossils
Fossils are known from: Aix-en-Provence, France (Late Oligocene), Baltic amber (Bartonian, Middle to Late Eocene), Barstow Formation, California, United States (Burdigalian to Langhian, Early to Middle Miocene), Bembridge Marls, Isle of Wight, UK (Priabonian, Late Eocene), Bitterfeld amber (Bartonian, Middle to Late Eocene), Brunn-Vösendorf, Austria (Late Miocene), Brunstatt, Haut-Rhin, France (Early Oligocene), Danish-Scandinavian amber (Bartonian, Middle to Late Eocene), Decín, Czech Republic (Early Miocene), Florissant, Colorado, United States (Late Eocene), Fushun amber, Liaoning, China (Ypresian, Early Eocene), Green River Formation, Colorado, United States (Lutetian, Middle Eocene), Joursac, Auvergne, France (Late Miocene), Montagne d'Andance, Saint-Bauzile, Ardèche, France (Early Turolian, Late Miocene), Oeningen, Switzerland (Messinian, Late Miocene), Radoboj, Croatia (Burdigalian, Early Miocene), Rovno amber (Priabonian, Late Eocene), Shanwang, China (Early Miocene).
Biology
Nests
There is great diversity of nest architecture: in ground, in dead wood, or structures constructed outside tree trunks and branches using carton (rigid material composed of masticated and moistened plant fibres) or silk.
Species belonging to three subgenera (Karavaievia in Asia; Dendromyrmex and Myrmobrachys in South America) use last-instar larvae to weave their nests, and incorporate sand particles, detritus and bits of plants into the silk sheet (e.g. Camponotus texens and Camponotus gombaki; Maschwitz et al. 1985). See also Camponotus nitidior, Camponotus chartifex, Camponotus senex
A twig-nesting species from Cairns, Queensland, Australia.
Association with Other Organisms
Species Uncertain
Coleoptera
Unknown species of Camponotus are prey for the tiger beetles Cicindela limbalis and Cicindela scutellaris lecontei in the United States (Valenti & Gaimari, 2000; Polidori et al., 2020).
Hymenoptera
Unspecified species within this genus are hosts for the following parasitic wasps (Universal Chalcidoidea Database):
Wasp Family | Wasp Species | Relationship |
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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 |
Orthoptera
An unknown species of Camponotus is a host for the cricket Myrmecophilus arboreus in New Guinea.
Fungi
Unknown species of Camponotus are hosts for the fungi Ophiocordyceps albacongiuae, Ophiocordyceps blakebarnesii, Ophiocordyceps rami and Ophiocordyceps septa (Araujo et al., 2018), and Ophiocordyceps irangiensis, Ophiocordyceps japonensis, Ophiocordyceps lloydii var. binata (Shrestha et al., 2017), and Torrubiella formicarum (in Ghana; Samson, et al., 1989).
All Associate Records for Genus
Taxon | Relationship | Associate Type | Associate Taxon | Associate Relationship | Locality | Source | Notes |
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Camponotus | host | cricket | Myrmecophilus arboreus | myrmecophile | New Guinea | ||
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 irangiensis | pathogen | Asia, North and South America | Shrestha et al., 2017 | |
Camponotus | host | fungus | Ophiocordyceps japonensis | pathogen | Asia, North and South America | Shrestha et al., 2017 | |
Camponotus | host | fungus | Ophiocordyceps lloydii var. binata | pathogen | Asia, North and South America | Shrestha et al., 2017 | |
Camponotus | host | fungus | Ophiocordyceps rami | pathogen | Shrestha et al. 2017; Araujo et al., 2018 | ||
Camponotus | host | fungus | Ophiocordyceps septa | pathogen | Shrestha et al. 2017; Araujo et al., 2018 | ||
Camponotus | host | fungus | Torrubiella formicarum | pathogen | Ghana | Samson, et al., 1989 | |
Camponotus | prey | tiger beetle | Cicindela limbalis | predator | United States | Valenti & Gaimari, 2000; Polidori et al., 2020 | |
Camponotus | prey | tiger beetle | Cicindela scutellaris lecontei | predator | United States | Valenti & Gaimari, 2000; Polidori et al., 2020 | |
Camponotus aberrans | host | fungus | Ophiocordyceps unilaterialis | pathogen | Africa | 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 | North America | Shrestha et al., 2017 | |
Camponotus angusticollis | prey | tiger beetle | Cicindela whithilli | predator | Western Ghats, India | Sinu et al., 2006 | |
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 | South America | Araujo et al., 2018; Shrestha et al., 2017; Sobczak et al., 2017 | |
Camponotus atriceps | host | fungus | Ophiocordyceps lloydii | pathogen | Colombia, Ecuador, Guyana | Araujo et al., 2018; Shrestha et al., 2017 | |
Camponotus atriceps | host | fungus | Ophiocordyceps unilaterialis | pathogen | North and South America | 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 | Brazil | Araujo et al., 2018; Evans et al., 2011; Shrestha et al., 2017 | |
Camponotus bispinosus | host | fungus | Ophiocordyceps camponoti-bispinosi | pathogen | South America | 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 | cricket | Myrmecophilus pergandei | myrmecophile | United States | ||
Camponotus castaneus | host | fungus | Ophiocordyceps kimflemingiae | pathogen | Araujo et al., 2018 | ||
Camponotus castaneus | host | fungus | Ophiocordyceps unilaterialis sensu lato | pathogen | Mangold et al., 2019 | ||
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 chromaiodes | host | cricket | Myrmecophilus pergandei | myrmecophile | United States | ||
Camponotus compressus | host | chalcid wasp | Epitranus chilkaensis | parasite | Narendran, 1989; Universal Chalcidoidea Database | primary host | |
Camponotus compressus | mutualist | butterfly | Lampides boeticus | Obregon et al. 2015 | |||
Camponotus crassus | host | strepsipteran | Stichotrema beckeri | parasite | Cook, 2019 | ||
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 | cricket | Myrmecophilus pergandei | myrmecophile | United States | ||
Camponotus herculeanus | host | eucharitid wasp | Stilbula cyniformis | parasite | Universal Chalcidoidea Database | primary host | |
Camponotus herculeanus | host | mite | Holostaspis flexuosa | parasite | Western Siberia, Russia | Joharchi et al., 2020 | ectoparasite |
Camponotus herculeanus | host | nematode | Oscheius dolichurus | parasite | Ontario | Nickle & Ayre, 1966 | |
Camponotus hippocrepis | host | fungus | Ophiocordyceps camponoti-hippocrepidis | pathogen | Araujo et al., 2018 | ||
Camponotus indianus | host | fungus | Ophiocordyceps camponoti-indiani | pathogen | Araujo et al., 2018; Shrestha et al., 2017 | ||
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 laevissimus | 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 | Brazil | Araujo et al., 2018; Evans et al., 2011; Shrestha et al., 2017 | |
Camponotus mitis | host | cricket | Myrmecophilus albicinctus | Myrmecophilus | India | ||
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 | cricket | Myrmecophilus pergandei | myrmecophile | United States | ||
Camponotus novaeboracensis | host | eucharitid wasp | Pseudochalcura gibbosa | parasite | Universal Chalcidoidea Database | primary host | |
Camponotus novogranadensis | host | fungus | Ophiocordyceps camponoti-novogranadensis | pathogen | Brazil | 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 parius | prey | tiger beetle | Cicindela duponti | predator | Western Ghats, India | Sinu et al., 2006 | |
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 planatus | host | strepsipteran | Caenocholax fenyesi | parasite | Cook, 2019 | ||
Camponotus punctatissimus | host | fungus | Ophiocordyceps unilaterialis | pathogen | Taiwan | Lin et al., 2020 | identification of host needs confirmation |
Camponotus punctulatus cruentus | host | strepsipteran | Caenocholax fenyesi | parasite | Cook, 2019 | ||
Camponotus punctulatus cruentus | host | strepsipteran | Myrmecolax ogloblini | parasite | Cook, 2019 | reported as "''Camponotus punctulatus cruentatus'' Forel" | |
Camponotus punctulatus cruentus | host | strepsipteran | Stichotrema beckeri | parasite | Cook, 2019 | ||
Camponotus renggeri | host | fungus | Ophiocordyceps camponoti-renggeri | pathogen | Araujo et al., 2018 | ||
Camponotus rufipes | host | fungus | Ophiocordyceps camponoti-rufipedis | pathogen | Brazil | Andersen et al., 2012; Araujo et al., 2018; Evans et al., 2011; Loreto et al., 2014; Shrestha et al., 2017 | |
Camponotus rufoglaucus | host | cricket | Myrmophilellus meneliki | myrmecophile | Ethiopia | Reichensperger, 1913 | |
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 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 | cricket | Myrmecophilus manni | myrmecophile | Mexico, United States | ||
Camponotus vicinus | host | nematode | Formicitylenchus oregonensis | parasite | Oregon, USA | Poinar, 2003 | |
Camponotus vividus | host | fungus | Ophiocordyceps unilaterialis | pathogen | Shrestha et al., 2017 |
Castes
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
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 Online Catalogue of the Ants of the World.
- 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:
Group I: Myrmopiromis
(Afrotropical and Malagasy regions, a few species from India)
1. Myrmopiromis
2. Myrmotrema
3. Myrmisolepis
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
2. Myrmostenus
4. Manniella
Group VII: Myrmobrachys
Subgroup transitional to Myrmaphaenus group
1. Myrmobrachys
3. Myrmomalis
4. Myrmeurynota
7. Myrmepomis
(Section by B. E. Boudinot, 19 February 2017.)
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