Temporal range: 53–0 Ma Ypresian, Early Eocene – Recent
|Formica ligniperda, now Camponotus ligniperda|
32 fossil 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 genus links included in this blurb for more detail.
- 1 Photo Gallery
- 2 Identification
- 3 Distribution
- 4 Biology
- 5 Castes
- 6 Morphology
- 7 Nomenclature
- 8 Common Names for Camponotus Subgenera
- 9 Subgenus Camponotus
- 10 Taxonomic History of Camponotus
- 11 Emery's Phylogenetic Hypothesis for Camponotus
- 12 References
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.
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
Emery's keys to the subgenera from Genera Insectorum
- 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 and Richness based on AntMaps
A twig-nesting species from Cairns, Queensland, Australia.
Unknown species of Camponotus are hosts for the fungi Ophiocordyceps albacongiuae, Ophiocordyceps blakebarnesii, Ophiocordyceps rami and Ophiocordyceps septa (Araujo et al., 2018).
• Antennal segment count 12 • Antennal club absent-gradual • Palp formula 6,4; 5,4 • Total dental count 4-9 • Spur formula 1 simple-barbulate, 1 simple-pectinate; 0, 0 • Eyes present • Scrobes absent • Caste polymorphic • Sting absent
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.
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")
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)
Group II: Myrmosaulus
(Malaysia, Australia, Madagascar)
Group III: Myrmamblys
Myrmopsamma subgroup (Africa)
Karavaievia subgroup (Indomalaya, etc.)
Mayria subgroup (Afrotropical, Malagasy)
2. Colobopsis [note: revived status as genus, Ward et al. (2015)]
Group IV: Tanaemyrmex
1. Tanaemyrmex [note: Emery's species groups may be particularly useful for understanding, and dividing, this probably "plesiotypic" taxon]
Group V: Myrmoxygenys group
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
2. Myrmaphaenus (New World)
Neotropical subgroup, transitional to Myrmobrachys group
Group VII: Myrmobrachys
Subgroup transitional to Myrmaphaenus group
(Section by B. E. Boudinot, 19 February 2017.)
- Agosti, D. 1991. Revision of the oriental ant genus Cladomyrma, with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae). Syst. Entomol. 16: 293-310 PDF (page 295, Camponotus in Formicinae, Formica genus group)
- Araújo, J.P.M., Evans, H.C., Kepler, R., Hughes, D.P. 2018. Zombie-ant fungi across continents: 15 new species and new combinations within Ophiocordyceps. I. Myrmecophilous hirsutelloid species. Studies in Mycology 90: 119–160 (DOI 10.1016/j.simyco.2017.12.002).
- Arnold, G. 1922. A monograph of the Formicidae of South Africa. Part V. Myrmicinae. Ann. S. Afr. Mus. 14: 579-674 (page 610, Camponotus in Camponotinae, Camponotini)
- Ashmead, W. H. 1905c. A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. Can. Entomol. 37: 381-384 (page 384, Camponotus in Camponotinae, Camponotini)
- Bingham, C. T. 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. London: Taylor and Francis, 506 pp. (page 347, Type-species: Formica ligniperda; by subsequent designation)
- Blaimer, B.B., Brady, S.G., Schultz, T.R., Lloyd, M.W., Fisher, B.L., & Ward, P.S. 2015. Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants. BMC Evolutionary Biology, 15:271 (DOI 10.1186/s12862-015-0552-5).
- Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp.
- Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 112, Camponotus in Formicinae, Camponotini, Camponotus as genus; page 113, Synopsis of subgenera [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)
- Bondroit, J. 1918. Les fourmis de France et de Belgique. Ann. Soc. Entomol. Fr. 87: 1-174 (page 65, Camponotus in Formicinae, Camponotini)
- Brady, S. G.; Schultz, T. R.; Fisher, B. L.; Ward, P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103:18172-18177.
- Brandão, C. R. F. 1991. Adendos ao catálogo abreviado das formigas da região Neotropical (Hymenoptera: Formicidae). Revista Brasileira de Entomologia 35:319-412.
- Brown, W. L., Jr. 1973b. A comparison of the Hylean and Congo-West African rain forest ant faunas. Pp. 161-185 in: Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Wash (page 179-185, All subgenera given as provisional junior synonyms of Camponotus (unconfirmed); see notes)
- Dalla Torre, K. W. von. 1893. Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann, 289 pp. (page 171, Camponotus in Camponotinae )
- Dlussky, G. M. 1997a. Genera of ants (Hymenoptera: Formicidae) from Baltic amber. Paleontol. Zh. 6: 50-62 (page 53, Camponotus (subgenus not given) senior synonym of Drymomyrmex)
- Duran-Ruiz, C. et al. 2013. Ants from the Miocene Totolapa amber (Chiapas, Mexico), with the first record of the genus Forelius (Hymenoptera, Formicidae). Can. J. Earth Sci. 50, 495-502 (dx.doi.org/10.1139/cjes-2012-0166).
- Emery, C. 1895l. Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 8: 685-778 (page 772, Camponotus in Camponotinae, Camponotini)
- Emery, C. 1896j. Saggio di un catalogo sistematico dei generi Camponotus, Polyrhachis e affini. Mem. R. Accad. Sci. Ist. Bologna (5)5:363-382 (page 376, combination in Calomyrmex)
- Emery, C. 1920b. Le genre Camponotus Mayr. Nouvel essai de la subdivision en sous-genres. Revue Zoologique Africaine (Brussels) 8:229-260.
- Emery, C. 1925d. Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum 183: 1-302 (page 59, 62, Synopsis of subgenera; Camponotus in Formicinae, Camponotini; page 35, Camponotus as subgenus)
- Emery, C.; Forel, A. 1879. Catalogue des Formicides d'Europe. Mitt. Schweiz. Entomol. Ges. 5: 441-481 (page 447, Camponotus in Camponotinae [Camponotidae])
- Forel, A. 1878c. Études myrmécologiques en 1878 (première partie) avec l'anatomie du gésier des fourmis. Bull. Soc. Vaudoise Sci. Nat. 15: 337-392 (page 367, Camponotus in Camponotinae [Camponotidae])
- Forel, A. 1886h. Études myrmécologiques en 1886. Ann. Soc. Entomol. Belg. 30: 131-215 (page 141, Camponotus in Camponotinae, Camponotini)
- Forel, A. 1892k. Les Formicides de l'Empire des Indes et de Ceylan. Part I. J. Bombay Nat. Hist. Soc. 7: 219-245 (page 223, Camponotus in Camponotinae [Camponotidae])
- Forel, A. 1893b. Sur la classification de la famille des Formicides, avec remarques synonymiques. Ann. Soc. Entomol. Belg. 37: 161-167 (page 165, Camponotus in Camponotinae, Camponotini)
- Forel, A. 1895b. A fauna das formigas do Brazil. Bol. Mus. Para. Hist. Nat. Ethnogr. 1: 89-139 (page 101, Camponotus in Camponotinae, Camponotini)
- Forel, A. 1899h. Formicidae. [part]. Biol. Cent.-Am. Hym. 3: 105-136 (page 130, Camponotus in Camponotinae, Camponotini)
- Forel, A. 1912j. Formicides néotropiques. Part VI. 5me sous-famille Camponotinae Forel. Mém. Soc. Entomol. Belg. 20: 59-92 (page 89, Camponotus in Camponotinae, Camponotini)
- Forel, A. 1914a. Le genre Camponotus Mayr et les genres voisins. Revue Suisse de Zoologie 22:257-276.
- Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 250, Camponotus in Camponotinae, Camponotini)
- Hong, Y.; Wu, J. 2000. The emendation of Shanwangella palaeoptera Zhang and its concerned problems. Geoscience (Beijing) 14: 15-20 (page 20, Camponotus (subgenus not given) senior synonym of Shanwangella)
- Kempf, W. W. 1972b. Catálogo abreviado das formigas da região Neotropical. Studia Entomologica 15:3-344.
- Mayr, G. 1861. Die europäischen Formiciden. Nach der analytischen Methode bearbeitet. Wien: C. Gerolds Sohn, 80 pp. (page 35, Camponotus in Formicinae [Formicidae])
- Mayr, G. 1862. Myrmecologische Studien. Verh. K-K. Zool.-Bot. Ges. Wien 12: 649-776 (page 652, Camponotus in Formicinae (in key) [Formicidae])
- Mayr, G. 1865. Formicidae. In: Reise der Österreichischen Fregatte "Novara" um die Erde in den Jahren 1857, 1858, 1859. Zoologischer Theil. Bd. II. Abt. 1. Wien: K. Gerold's Sohn, 119 pp. (page 6, Camponotus in Formicinae [Formicidae])
- Mayr, G. 1868c. Die Ameisen des baltischen Bernsteins. Beitr. Naturkd. Preuss. 1: 1-102 (page 26, Camponotus in Formicinae [Formicidae])
- Moreau, C. S.; Bell, C. D.; Vila, R.; Archibald, S. B.; Pierce, N. E. 2006. Phylogeny of the ants: diversification in the age of angiosperms. Science (Washington, D. C.) 312:101-104.
- Santschi, F. 1921f. Retouches aux sous-genres de Camponotus. Ann. Soc. Entomol. Belg. 61: 310-312 (page 310, Myrmisolepis as subgenus of Camponotus)
- Santschi, F. 1926c. Nouvelles notes sur les Campontous. Rev. Suisse Zool. 33: 597-618 (page 604, Paramyrmambly as subgenus of Camponotus)
- Santschi, F. 1928e. Nouvelles fourmis d'Australie. Bull. Soc. Vaudoise Sci. Nat. 56: 465-483 (page 483, Thlipsepinotus as subgenus of Camponotus)
- Shattuck, S. O. 1999. Australian ants. Their biology and identification. Collingwood, Victoria: CSIRO Publishing, xi + 226 pp.
- Smith, F. 1871a. A catalogue of the Aculeate Hymenoptera and Ichneumonidae of India and the Eastern Archipelago. With introductory remarks by A. R. Wallace. [part]. J. Linn. Soc. Lond. Zool. 11: 285-348 (page 306, Camponotus as subgenus of Formica)
- Taylor, R. W.; Brown, D. R. 1985. Formicoidea. Zoological Catalogue of Australia 2:1-149, 306-348.
- Ward, P.S., Blaimer, B.B., Fisher, B.L. 2016. A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinomyrmex. Zootaxa 4072 (3): 343–357 (doi 10.11646/zootaxa.4072.3.4).
- Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 144, Camponotus in Camponotinae, Camponotini)
- Wheeler, W. M. 1913a. Corrections and additions to "List of type species of the genera and subgenera of Formicidae". Annals of the New York Academy of Sciences 23:77-83.
- Wheeler, W. M. 1915i . The ants of the Baltic Amber. Schr. Phys.-Ökon. Ges. Königsb. 55: 1-142 (page 138, Camponotus in Camponotinae, Camponotini)
- Wheeler, W. M. 1922i. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bull. Am. Mus. Nat. Hist. 45: 631-710 (page 703, Synopsis of subgenera, Camponotus in Formicinae, Camponotini)