Camponotus

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Camponotus
Temporal range: 53–0 Ma 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
Diversity
1,504 species
1 fossil species
(Species Checklist, Species by Country)

Camponotus ligniperda casent0173649 profile 1.jpg

Camponotus ligniperda

Camponotus ligniperda casent0173649 dorsal 1.jpg

Specimen Label

Synonyms

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).

Taxonomically, Camponotus and the tribe Camponotini more broadly is one of the most challenging puzzles in the Formicidae, and not merely due to the massive size of these taxa (1,504 species and subspecies in Camponotus). Although some genera in the tribe are reasonably identifiable based on external morphology (e.g., Ward et al. 2016), others, such as the fundamental distinction between Colobopsis—which is sister to all other Camponotini—and the hyperdiverse Camponotus is challenging even with extant material in hand and under the microscope (Ward and Boudinot, 2021; Boudinot et al., 2024).


Photo Gallery

  • Camponotus Shattuck 53201 Danum Valley Sabah-web.jpg
  • Camponotus Shattuck 54238 Danum Valley Sabah-web.jpg
  • Camponotus pennsylvanicus feeding on a droplet of honey.
  • Shattuck C16190-web, ANIC32-030620, Camponotus, Cairns, Qld.jpg
  • Camponotus Shattuck 54662 Danum Valley Sabah-web.jpg
  • A queen from Dominican amber.
  • Mermithid nematode exposed in the gaster of Camponotus sp. from the Sierra Nevada, California (Poinar, 2012, Fig. 2).
  • A Camponotus sp. infected by the fungus Ophiocordyceps unilateralis from San Pedrillo, Puntarenas, Costa Rica. Photo by Katja Schulz.
  • Camponotus compressus minor worker carrying a termite dealate. Dehradun, Uttarakhand, India. Photo by Yathumon M A.

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.

Distinguishing Colobopsis from Camponotus

To differentiate Colobopsis from Camponotus Ward et al. (2016) proposed the following global key:

1 Not occurring in Fiji or New Caledonia ... 2

- Occurring in Fiji ... 3

- Occurring New Caledonia ... 4


2 Generally small species, HW 0.65–1.10 (except cylindrica-group of Southeast Asia with HW 1.20–1.70, and facies as in Figures 4 and 5); either antennal insertions relatively well separated, such that ASM/HW 0.36–0.47 and ASM/CLW 0.66–0.98, and/or clypeus relatively narrow, such that CLW/CLL 0.96–1.32; antennal insertions occurring at about midlength of frontal carinae; anterolateral extremities of clypeus set off from rest of clypeus by a sulcus or impression, so clypeus appears to lack prominent anterolateral extensions (Figures 2–5) ... Colobopsis

- Small to large species, HW 0.70–3.00; antennal insertions less well separated, such that ASM/HW 0.22–0.35 and ASM/CLW 0.35–0.68; clypeus variable in shape but in smaller species with HW 0.70–1.35 (e.g., Camponotus (Myrmamblys), C. (Myrmentoma) and C. (Pseudocolobopsis)) clypeus tending to be relatively broad, such that CLW/CLL 1.25–1.62, although exceptions occur (e.g., in some C. (Pseudocolobopsis) species) (Figures 14–15); antennal insertions usually occurring in front of midlength of frontal carinae; clypeus typically with prominent anterolateral extensions (Figure 15) ... Camponotus


3 With conspicuous long setae, gracile legs, and a shield-shaped clypeus with prominent anterolateral extensions (Figure 16) ... Camponotus chloroticus

- Without the combination of conspicuous long setae and gracile legs; clypeus lacking prominent anterolateral extensions (Figures 8–11) ... Colobopsis


4 Small species, HW 0.68–1.04; antennal insertions more widely separated (ASM/HW 0.34–0.39 and ASM/CLW 0.64–0.77) (Figures 18–19); clypeus tending to be less broad (CLW/CLL 1.15–1.40) ... Colobopsis

- Small to medium-sized species, HW 0.75–2.10; antennal insertions less well separated (ASM/HW 0.25–0.29 and ASM/CLW 0.46–0.55); clypeus varying in shape, but if HW < 1.05 (e.g., Camponotus pulchellus complex) (Figure 17) then clypeus tending to be broader (CLW/CLL 1.25–1.60) ... Camponotus


AntWeb icon 02.png See images of species within this genus

Keys including this Genus

Keys to Subgenera or Species Groups in this Genus

Keys to Species in this Genus

Emery's keys to the subgenera from Genera Insectorum

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

Species Groups

Distribution

Distribution and Richness based on AntMaps

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 308 145 203 113 50 477 127 239
Total Species 2851 1736 3047 932 840 4391 1767 2925

Fossils

Fossils are known from: Aix-en-Provence, France (Late Oligocene), Baltic amber, Baltic Sea region, Europe (Priabonian, 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, Baltic Sea region, Europe (Priabonian, Late Eocene), Brunn-Vösendorf, Austria (Late Miocene), Brunstatt, Haut-Rhin, France (Early Oligocene), Canyon Ferry Reservoir, Montana, United States (Rupelian, Oligocene) (an unidentified species, LaPolla, 2023), Danish-Scandinavian amber (Priabonian, Late Eocene), Decín, Czechia (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, Baltic Sea region, Europe (Priabonian, Late Eocene), Shanwang, China (Early Miocene), Zhangpu amber, Zhangpu County, Fujian Province, China (Miocene) (an unidentified species, Wang et al., 2021).

Takahashi & Aiba (2023) comment on a winged Camponotus fossils from the Chibanian (Middle Pleistocene) Shiobara Group, Tochigi Prefecture, Japan. However, B. Boudinot (03.Nov.2023) notes that these specimens are misidentified to genus; while it is plausible that they may be Formicinae, it is not immediately obvious to which genus they should be attributed.

Biology

Nesting Habits

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.

Shattuck C16190-web, ANIC32-030620, Camponotus, Cairns, Qld.jpg Shattuck C16415-1, ANIC32-030620, Camponotus, Cairns, Qld.jpg Shattuck C16458-1, ANIC32-030620, Camponotus, Cairns, Qld.jpg Shattuck C16469-1, ANIC32-030620, Camponotus, Cairns, Qld.jpg

Ant Gardens

Four unidentified species of Camponotus from peninsular Malaysia are known to form ant-gardens (i.e., they are able to initiate ant gardens or are restricted to ant gardens) (Kaufmann at al., 2001; Kaufmann, 2002 (noted as ant-garden initiator); Kaufmann & Maschwitz, 2006; Orivel & Leroy, 2011).

Association with Other Organisms

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Unknown species of Camponotus are involved in the following associations:

Associate Type Associate Taxon Relationship Locality Source Notes
Braconid wasp Elasmosoma berolinense Parasitoid Quevillon, 2018 encounter mode primary; direct transmission; transmission outside nest
Encyrtid wasp Arketypon vaderi Parasitoid Quevillon, 2018
Encyrtid wasp Arketypon vaderi Parasitoid Universal Chalcidoidea Database associate
Eucharitid wasp Ancylotropus manipurensis Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Lophyrocera variabilis Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Mateucharis rugulosa Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Pseudochalcura americana Parasitoid Quevillon, 2018 encounter mode secondary; direct transmission; transmission outside nest
Eucharitid wasp Pseudochalcura nigrocyanea Parasitoid Quevillon, 2018 encounter mode secondary; direct transmission; transmission outside nest
Eucharitid wasp Rhipipalloidea madangensis Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Rhipipalloidea sp. Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Stilbula cyniformis Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Stilbuloida doddi Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Zulucharis campbelli Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eucharitid wasp Ancylotropus manipurensis Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Lophyrocera variabilis Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Mateucharis rugulosa Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Pseudochalcura gibbosa Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Pseudochalcura nigrocyanea Parasitoid Universal Chalcidoidea Database associate
Eucharitid wasp Rhipipalloidea sp. Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Rhipipalloidea madangensis Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Stilbula cyniformis Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Stilbuloida doddi Parasitoid Universal Chalcidoidea Database primary host
Eucharitid wasp Zulucharis campbelli Parasitoid Universal Chalcidoidea Database primary host
Eulophid wasp Horismenus microdonophagus Parasitoid Quevillon, 2018
Eulophid wasp Horismenus myrmecophagus Parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Eulophid wasp Horismenus microdonophagus Parasitoid Universal Chalcidoidea Database associate
Eulophid wasp Horismenus myrmecophagus Parasitoid Universal Chalcidoidea Database primary host
Eulophid wasp Pediobius acraconae Parasitoid Quevillon, 2018
Eurytomid wasp Camponotophilus delvarei Parasitoid Universal Chalcidoidea Database associate
Phorid fly Apocephalus coquilletti Parasitoid Quevillon, 2018
Phorid fly Apocephalus crucicauda Parasitoid Quevillon, 2018
Phorid fly Apocephalus crucicauda Parasitoid phorid.net attacked
Phorid fly Apocephalus similis Parasitoid Quevillon, 2018
Phorid fly Apocephalus similis Parasitoid phorid.net attacked
Phorid fly Diocophora appretiata Parasitoid Quevillon, 2018
Phorid fly Diocophora disparifrons Parasitoid Quevillon, 2018
Phorid fly Rhyncophoromyia gymnopleura Parasitoid Quevillon, 2018
Microdon fly Microdon piperi Parasitoid Quevillon, 2018
Cricket Myrmecophilus arboreus Myrmecophile New Guinea
Cricket Myrmecophilus baronii Myrmecophile Mediterranean region Stalling & Cassar, 2020
Tiger beetle Cicindela limbalis Predator United States Valenti & Gaimari, 2000; Polidori et al., 2020
Tiger beetle Cicindela scutellaris lecontei Predator Valenti & Gaimari, 2000; Polidori et al., 2020
Nematode Steinerema carpocapsae Parasitoid Quevillon, 2018
Strepsipteran Apocephalus conecitonis Parasitoid Quevillon, 2018
Strepsipteran Apocephalus paraponerae Parasitoid Quevillon, 2018
Fungus Beauveria bassiana Parasitoid Quevillon, 2018
Fungus Gibellula formicarium Parasitoid Quevillon, 2018
Fungus Laboulbenia camponoti Parasitoid Quevillon, 2018
Fungus Ophiocordyceps acerosa Parasitoid Quevillon, 2018
Fungus Ophiocordyceps albacongiuae Parasitoid Quevillon, 2018
Fungus Ophiocordyceps albacongiuae Parasitoid Araujo et al., 2018
Fungus Ophiocordyceps blakebarnesii Parasitoid Quevillon, 2018
Fungus Ophiocordyceps blakebarnesii Parasitoid Araujo et al., 2018
Fungus Ophiocordyceps irangiensis Parasitoid Araujo et al., 2018
Fungus Ophiocordyceps japonensis Parasitoid Asia, North and South America Shrestha et al., 2017
Fungus Ophiocordyceps lloydii binata Parasitoid Asia, North and South America Shrestha et al., 2017
Fungus Ophiocordyceps lloydii binata Parasitoid Quevillon, 2018
Fungus Ophiocordyceps rami Parasitoid Quevillon, 2018
Fungus Ophiocordyceps rami Parasitoid Shrestha et al. 2017; Araujo et al., 2018
Fungus Ophiocordyceps septa Parasitoid Shrestha et al. 2017; Araujo et al., 2018
Fungus Ophiocordyceps septa Parasitoid Quevillon, 2018
Fungus Ophiocordyceps unilateralis Parasitoid Quevillon, 2018
Fungus Stilbella burmensis Parasitoid Quevillon, 2018
Fungus Torrubiella formicarum Parasitoid Ghana Samson, et al., 1989

All Associate Records for Genus

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Flight Period

All Flight Records for Genus

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Life History Traits

  • Mean colony size: 1000-100000+ (Greer et al., 2021)
  • Compound colony type: inquilinism (Greer et al., 2021)
  • Nest site: hypogaeic; arboreal (Greer et al., 2021)
  • Diet class: omnivore (Greer et al., 2021)
  • Foraging stratum: subterranean/leaf litter; arboreal (Greer et al., 2021)
  • Foraging behaviour: cooperative (Greer et al., 2021)

Castes

Morphology

Worker Morphology

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• 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: >100 ommatidia • Scrobes: absent • Pronotal Spines: absent • Mesonotal Spines: absent • Propodeal Spines: absent • Petiolar Spines: absent • Caste: polymorphic • Sting: absent • Metaplural Gland: absent; present • Cocoon: present


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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Phylogeny

Formicinae
Myrmelachistini
Lasiini
Melophorini
Formicini
Gesomyrmecini

Gesomyrmex  (7 species, 12 fossil species)

Oecophyllini

Oecophylla  (15 species, 16 fossil species)

Plagiolepidini
Gigantiopini

Gigantiops  (1 species, 0 fossil species)

Santschiellini

Santschiella  (1 species, 0 fossil species)

Myrmoteratini

Myrmoteras  (42 species, 0 fossil species)

Camponotini
⊟

See Phylogeny of Formicinae for details.

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, 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.]
  • DOLOPHRA [junior synonym of Camponotus]
    • Dolophra Wu, J. & Wang, 1994: 35. Type-species: Dolophra politae, by original designation.
    • Dolophra junior synonym of Camponotus: Bolton, 1995b: 27.
    • Dolophra junior synonym of Camponotus (Colobopsis): Bolton, 2003: 113, 268.
    • Dolophra junior synonym of Camponotus: Ward & Boudinot, 2021: 44.
  • †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.)

References

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