Belongs within: Camponotini.
Contains: Camponotus (Camponotus), Camponotus (Myrmamblys), Camponotus (Tanaemyrmex).
Camponotus is a diverse and cosmopolitan genus of ants, generally with a characteristic rounded profile to the rear of the mesosoma. They are commonly referred to as carpenter ants owing to the habit of some species of building nests in galleries in dead wood; however, not all species in the genus exhibit this behaviour. Workers of most Camponotus species are polymorphic, with distinct major and minor forms present (some species may also possess media forms). Major workers typically have a much more massive head than minors, with larger mandibles.
Species have been divided between several subgenera but many of the resultant groupings are poorly defined and require formal phylogenetic testing; as a result, many authors eschew their usage. Phasmomyrmex, a small group of Afrotropical species with rectangular or trapezoidal heads and monomorphic workers, has historically been treated as a distinct genus but was reclassified as a subgenus of Camponotus by Ward et al. (2016). Pseudocolobopsis, the false cork-head ants, is a Neotropical group of which the majors and queens have an elongate, often anteriorly obliquely truncate head. Forelophilus is a group of small, blackish ants found in south-east Asia with a strong transverse ridge across the propodeum.
Camponotus: a sugary high
Published 25 March 2017
I think I may have said before that Australia is the land of ants. When travelling in Australia’s arid regions (i.e. most of the continent), ants are often the most visible animals about. Perhaps the most visible of all Australia’s ants are the meat ants (Iridomyrmex), but not too far behind them are the sugar ants of the genus Camponotus.
Camponotus is a genus of the ant subfamily Formicinae found pretty much everywhere around the world that ants are to be found. It is massively diverse: well over 1000 species have been assigned to this genus over the years, with probably more to be described. They are correspondingly diverse in habits and appearance. Some are among the giants of the ant world, others are much smaller. Some form massive colonies that are difficult to miss and forage during the day, others are more retiring and emerge only at night. Some construct their nests in holes under the grounds, others hollow out wood or use the holes left by other wood-boring insects. Most (but not all) Camponotus species exhibit some form of worker polymorphism: rather than having just a single worker caste, a colony will often include large major workers and much smaller minor workers, with the two forms superficially looking quite different. Sometimes the distinction between majors and minors will be quite clear, other times there will also be workers of intermediate sizes. In some Australian species, known as honeypot ants, there are specialised workers called ‘repletes’ who spend their lives hanging in one spot inside the nest, being fed by the other active workers until their gasters swell into engorged round balls. These repletes serve the colony as a living larder, able to regurgitate their stored excess of food when needed by their nestmates. Despite all this diversity, most Camponotus species are readily recognisable as Camponotus: they usually lack spines on the mesosoma (the ‘thorax’), the back end of which is narrow and often arched. This smoothness and slimness gives Camponotus a distinctive look that kind of puts me in mind of the ant version of a greyhound. The majority of Camponotus species also differ from other ants in lacking the metapleural gland, a gland producing an antibiotic chemical whose opening is usually visible near the rear of the mesosoma.
Camponotus species have been referred to in Australia as ‘sugar ants’ in reference to their diet, which is commonly dominated by the sugary excretions of plant-sucking bugs that they attend. In other parts of the world, they have sometimes been referred to as ‘carpenter ants’ in reference to the wood-tunneling habits of their most notorious representatives. Bug-derived honeydew is high in sugar but low in other essential nutrients, so the ants also feed on things such as the scavenged bodies of the bugs themselves after death. They are also probably assisted in meeting their nutritive needs by Blochmannia, an endosymbiotic bacterium that infests specialised cells in the gut of Camponotus and closely related genera (Wernegreen et al. 2009). Genetic data from the endosymbiont indicates that it probably synthesises nutrients the ant does not otherwise ingest. It may also play some role in compensating for an absence of metapleural gland secretions. As well as the gut, Blochmannia infest the ovaries of reproductive females and are passed to the next generation via the developing oocytes. Phylogenetic analysis of Blochmannia indicates that it is closely related to other endosymbiotic bacteria found in mealybugs, and it is possible that the ancestors of Camponotus picked it up in the course of feeding on honeydew.
The sheer size of Camponotus as a genus has been a challenge to understanding relationships within the genus. Over thirty subgenera have been proposed at one time or another, but many of these are poorly defined and many authors eschew using them in favour of informal species groups. It does not help matters that, since the early 20th century, most reviews of Camponotus have been conducted at a local rather than a global level. Those studies that have touched on Camponotus phylogeny in recent years suggest the need for a large-scale revision, with few of the subgenera supported as monophyletic.
Systematics of Camponotus
Characters (from Heterick 2009): Antenna with twelve segments (including scape); propodeal angle (if present) rounded; metapleural gland opening inconspicuous, without fringe of long setae; first gastral tergite much less than half total length of gaster; major and minor worker castes, at least, always present, media workers often present.
Camponotus Mayr 1861T99 [incl. Myrmophyma Forel 1912TB85, Thlipsepinotus Santschi 1928TB85]
|--+--C. gibbinotus Forel 1902WBF16, TB85
| `--C. (Phasmomyrmex) [incl. Myrmacantha, Myrmorhachis]WBF16
| |--C. (P.) buchneri [incl. *Phasmomyrmex sericeus]WBF16
| | |--C. b. buchneriWBF16
| | `--C. b. camerounensis Ward, Blaimer & Fisher 2016 [=Phasmomyrmex buchneri griseus (preoc.)]WBF16
| |--C. (P.) aberransWBF16
| |--C. (P.) paradoxusWBF16
| | |--C. p. paradoxusWBF16
| | `--C. p. cupreusWBF16
| `--C. (P.) wolfiWBF16
`--+--C. (Pseudocolobopsis)WBF16
| |--C. (P.) claviscapusWBF16
| `--C. (P.) cordincola [=C. (Colobopsis) cordincola]WBF16
`--+--+--C. hyattiWBF16
| `--C. maritimusWBF16
`--+--C. bedotiWBF16
`--C. (Forelophilus)WBF16
|--C. (F.) javaensis Ward, Blaimer & Fisher 2016 [=*Forelophilus overbecki (preoc.)]WBF16
|--C. (F.) philippinensisWBF16
`--C. (F.) stefanschoedliWBF16
Camponotus incertae sedis:
C. abdominalisCM07
C. adami Forel 1910TB85
C. aeneopilosus Mayr 1862TB85
|--C. a. aeneopilosusTB85
`--C. a. flavidopubescens Forel 1902TB85
C. afflatus Viehmeyer 1925 [=C. (Myrmosaga) afflatus]TB85
C. albivillosus Zhou 2001Z01
C. angusticollis (Jerdon 1851) (see below for synonymy)B03
C. arcuatus Mayr 1876TB85
|--C. a. arcuatusTB85
`--C. a. aesopus Forel 1907TB85
C. armstrongi McAreavey 1949TB85 [=C. (Myrmogonia) armstrongiWBF16]
C. auratiacus Zhou 2001Z01
C. auriventris Emery 1889B03
C. aurocinctus (Smith 1858) [=Formica aurocincta]TB85
C. baldacciiES12
C. barbatus Roger 1863B03
C. basalis Smith 1878 [incl. C. maculatus lobinieri Forel 1902]B03
C. bifossus [=C. (Colobopsis) bifossus]WBF16
C. bigenus Santschi 1919 [=C. (Myrmocamelus) bigenus]TB85
C. binghami Forel 1894B03
C. branneriP92
C. breviscapus Zhou 2001Z01
C. buddhae Forel 1892B03
C. cameratus Viehmeyer 1925 [=C. (Myrmogonia) cameratus]TB85
C. capito Mayr 1876 [=C. (*Myrmophyma) capito]TB85
|--C. c. capitoTB85
`--C. c. ebeninithorax Forel 1915 [=C. (Myrmophyma) capito ebeninithorax]TB85
C. carbonariusB03
C. carin Emery 1889 [=C. dorycus carin]B03
C. ceriseipes Clark 1938 [=C. (Myrmophyma) ceriseipes]TB85
C. chalceoides Clark 1938 [=C. (Myrmophyma) chalceoides]TB85
C. chalceus Crawley 1915 [=C. (Myrmosaga) chalceus]TB85
C. cinereus Mayr 1876TB85
|--C. c. cinereusTB85
|--C. c. amperei Forel 1913 [=C. (Myrmocamelus) cinereus amperei]TB85
`--C. c. notterae Forel 1907TB85
C. clarior Forel 1902H09, TB85 [=C. nigriceps clariorTB85]
C. claripes Mayr 1876 [=C. (*Thlipsepinotus) claripes]TB85
|--C. c. claripesTB85
|--C. c. elegans Forel 1902TB85
|--C. c. inverallensis Forel 1910TB85
|--C. c. marcens Forel 1907TB85
|--C. c. minimus Crawley 1922 [=C. (Myrmophyma) claripes minima]TB85
|--C. c. nudimalis Forel 1913TB85
|--C. c. orbiculatopunctatus Viehmeyer 1925 [=C. (Myrmophyma) claripes orbiculatopunctatus]TB85
`--C. c. piperatus Wheeler 1933 [=C. (Myrmophyma) claripes piperatus]TB85
C. compressus (Fabr. 1787)B03 [=Formica compressaB03, C. maculatus compressusB03, C. rubripes compressusE89]
C. confucii Forel 1894B03
C. consectator (Smith 1858) [=Formica consectator]TB85
C. consobrinus (Erichson 1842) [=Formica consobrina; incl. C. dimidiatus Roger 1863]TB85
C. cowlei Froggatt 1896TB85
C. coxalisE89
C. crassisquamis Forel 1902B03
C. crassusMC05
C. crenatus Mayr 1876TB85
C. cruentatus (Latreille 1802) [=Formica cruentata]TB85
|--C. c. cruentatusTB85
`--C. c. aspera Menozzi 1925 [=C. (Myrmosericus) cruentatus aspera]TB85
C. darlingtoni Wheeler 1934H09, TB85 [=C. (Myrmophyma) darlingtoniTB85, C. (M.) rottnesti Donisthorpe 1941TB85]
C. denticulatus Kirby 1896TB85
C. dichrous Forel 1879 [=C. maculatus dichrous; incl. C. dichrous var. kattensis]B03
C. discors Forel 1902 [=C. maculatus discors]TB85
|--C. d. discorsH09
|--C. d. angustinodus (n. d.)H09
`--C. d. yarrabahensis Forel 1915 [=C. (Myrmoturba) discors yarrabahensis]TB85
C. dolendus Forel 1892 [=C. rufoglaucus r. dolendus]Z01
C. donnellaniH09
C. dorycus (Smith 1860) [=Formica dorycus]TB85
|--C. d. dorycusTB85
`--C. d. confusus Emery 1887TB85
C. dromas Santschi 1919 [=C. (Myrmocamelus) dromas]TB85
C. dryandraeH09
C. eremicus Wheeler 1915 [=C. (Myrmogonia) eremicus]TB85
C. erythropus Viehmeyer 1925 [=C. (Myrmosaga) erythropus]TB85
C. esau Forel 1915 [=C. (Myrmocamelus) esau]TB85
C. eugeniaeA61
C. evae Forel 1910TB85
|--C. e. evaeTB85
`--C. e. zeuxis Forel 1915 [=C. (Myrmogonia) evae zeuxis]TB85
C. exasperatusE89
C. extensus Mayr 1876TB85
C. festinus (Smith 1857) [=Formica festina]B03
C. fictor Forel 1902 [=C. (Colobopsis) fictor]TB85
|--C. f. fictorTB85
`--C. f. augustulus Viehmeyer 1925 [=C. (Colobopsis) fictor augustulus]TB85
C. fieldeae Forel 1902TB85
C. fieldellus Forel 1910TB85
C. floridanusPK17
C. fornaronis Forel 1892 [=C. mistura fornaronis]B03
C. froggatti Forel 1902TB85
C. fuscivillosusZ01
C. gouldianus Forel 1922TB85
C. gretae Forel 1902B03
C. habereiT99
C. hartogi Forel 1902 [incl. C. (Myrmosaga) ferruginipes Crawley in Poulton & Crawley 1922]TB85
C. herculeanusMS01
C. holosericeus Emery 1889B03
C. horni Kirby 1896TB85
C. inflatus Lubbock 1880 [incl. C. (Myrmamblys) aurofasciatus Wheeler 1915]TB85
C. infuscus Forel 1892 [=C. maculatus infuscus]B03
C. innexus Forel 1902TB85
C. insipidus Forel 1893 (n. d.)TB85
C. integer [=C. sericeus var. integer]B03
C. intrepidus (Kirby 1818) [=Formica intrepida]TB85
|--C. i. intrepidus [incl. Formica agilis Smith 1858, C. magnus Mayr 1862]TB85
`--C. i. bellicosus Forel 1902TB85
C. invidus Forel 1892B03
C. irritans (Smith 1857) [=Formica irritans, C. maculatus irritans; incl. C. agnatus Roger 1863]B03
C. janeti Forel 1895TB85
C. janforrestaeWBF16
C. jejuensisT99
C. jianghuaensis Xiao & Wang 1989Z01
C. johnclarki [=Notostigma johnclarki]H09
C. laevigatusWB03
C. lamarcki Forel 1892B03
C. largicepsZ01
C. lasiselene Wang & Wu 1994Z01
C. latrunculus Wheeler 1915 [=C. (Myrmoturba) latrunculus]TB85
|--C. l. latrunculusTB85
`--C. l. victoriensis Santschi 1928 [=C. (Myrmoturba) latrunculus victoriensis]TB85
C. leae Wheeler 1915 [=C. (Myrmosphincta) leae]TB85
C. lespesiMC05
C. lividicoxis Viehmeyer 1925 [=C. (Myrmophyma) lividicoxis]TB85
C. longideclivisH09
C. longifaciesH09
C. lownei Forel 1895 [=Formica nitida Lowne 1865 non Smith 1858]TB85
C. luteus (Smith 1858) [=Formica lutea]B03
C. macareaveyi [=C. (Myrmogonia) macareaveyi]WBF16
C. mackayensis Forel 1902FT08, TB85 [=C. reticulatus mackayensisTB85]
C. marginatus (Latr. 1798)B03 (see below for synonymy)
C. melichloros Kirby 1888FT08
C. melleus (Say 1835)SW07
C. mendax Forel 1895 [=C. sericeus var. mendax]B03
C. micansE89
C. michaelseni Forel 1907TB85 [incl. C. walkeri bardus Forel 1910H09, TB85]
C. midas Froggatt 1896TB85
C. minus Wang & Wu 1994Z01
C. mistura (Smith 1857) [=Formica mistura]B03
C. mitis (Smith 1858)B03 (see below for synonymy)
C. molossus Forel 1907TB85
C. myoporus Clark 1938 [=C. (Tanaemyrmex) myoporus]TB85
C. nicobarensis Mayr 1865 (see below for synonymy)B03
C. nigriceps (Smith 1858) [=Formica nigriceps]TB85
|--C. n. nigricepsTB85
|--C. n. lividipes Emery 1887TB85
|--C. n. obniger Forel 1902TB85
`--C. n. pallidiceps Emery 1887TB85
C. nigroaeneus (Smith 1858) [=Formica nigroaenea]TB85
|--C. n. nigroaeneusTB85
`--C. n. divus Forel 1907TB85
C. nirvanae Forel 1893B03
C. nitidiceps Viehmeyer 1925 [=C. (Myrmophyma) nitidiceps]TB85
C. oblongus (Smith 1858) [=Formica oblonga]B03
C. oetkeri Forel 1910TB85
|--C. o. oetkeriTB85
`--C. o. voltai Forel 1913TB85 [=C. (Myrmogonia) oetkeri voltaiTB85; incl. C. rudisH09]
C. oxleyi Forel 1902TB85
C. pallensE89
C. pallidus (Smith 1857) [=Formica pallida; incl. C. rubripes subnudus Emery 1889]B03
C. papuaK91
C. paria Emery 1889 [=C. micans paria, C. rufoglaucus paria]B03
C. pawseyiH09
C. pellax Santschi 1919 [=C. (Myrmocamelus) pellax]TB85
C. perjurusH09
C. pilicornisES12
C. pitjantjataraeH09
C. postcornutus Clark 1930 [=C. (Tanaemyrmex) postcornutus]TB85
C. prosseriH09
C. prostansH09
C. pseudoirritans Wu & Wang 1989Z01
C. punctiventris Emery 1920 [=C. (Myrmogonia) punctiventris]TB85
C. radiatus Forel 1892B03
C. renggeriB14
C. reticulatus Roger 1863FT08 [incl. C. reticulatus var. latitansB03]
C. rubiginosus Mayr 1876TB85
C. rubripesG89
|--C. r. rubripesG89
`--C. r. cognatusG89
C. rufoglaucus (Jerdon 1851) [=Formica rufoglauca; incl. C. redtenbacheri Mayr 1862]B03
C. rufus Crawley 1925 [=C. (Dinomyrmex) rufus]TB85
C. rugipesWL04
C. ‘sanguinea’ McAreavey 1949 non C. japonicus var. sanguinea Karavajew 1929 [=C. (Myrmogonia) sanguinea]TB85
C. scottiH09
C. scratius Forel 1907TB85
|--C. s. scratiusTB85
`--C. s. nuntius Forel 1907TB85
C. semicarinatus (Forel 1895) [=Colobopsis rufifrons semicarinatus]TB85
C. sericeiventrisB10
C. sericeus (Fabr. 1798) (see below for synonymy)B03
C. siemsseni Forel 1901B03
C. simpsoni McArthur 2003H09
C. simulator Forel 1915 [=C. (Dinomyrmex) simulator]TB85
C. spanis Xiao & Wang 1989Z01
C. spenceri Clark 1930 [=C. (Tanaemyrmex) spenceri, C. reticulatus Kirby 1896 non Roger 1863]TB85
C. spinitarsus Emery 1920 [=C. (Dinomyrmex) spinitarsus]TB85
C. sponsorum Forel 1910TB85
C. subnitidus Mayr 1876TB85
|--C. s. subnitidusTB85
|--C. s. famelicus Emery 1887TB85
`--C. s. longinodis Forel 1915 [=C. (Dinomyrmex) subnitidus longinodis]TB85
C. suffusus (Smith 1858) [=Formica suffusa]TB85
|--C. s. suffusus [incl. Formica piliventris Smith 1858, C. schencki Mayr 1862]TB85
`--C. s. bendigensis Forel 1902TB85
C. sylvaticusES12
C. tasmani Forel 1902TB85
C. taylori Forel 1892 [=C. maculatus taylori; incl. C. taylori var. infuscoides]B03
C. terebrans (Lowne 1865)H09, TB85 [=Formica terebransTB85]
C. testaceipes (Smith 1858) [=Formica testaceipes]TB85
C. thraso Forel 1893 [=C. maculatus thraso]B03
C. tinctus (Smith 1858)E14, B03 [=Formica tinctaB03]
C. tricoloratus Clark 1941 [=C. (Tanaemyrmex) tricoloratus]TB85
C. tristis Clark 1930 [=C. (Myrmophyma) tristis]TB85
C. tritschleri [=C. (Colobopsis) tritschleri]WBF16
C. tumidus Crawley 1922 [=C. (Myrmogonia) tumidus]TB85
C. universitatisES12
C. varians Roger 1863B03
C. variegatus (Smith 1858) [=Formica variegata; incl. C. (Myrmoturba) variegatus hawaiensis Emery 1920]Z01
C. versicolor Clark 1930 [=C. (Myrmosaulus) versicolor]TB85
C. villosus Crawley 1915 [=C. (Myrmoturba) villosa]TB85
C. walkeri Forel 1893TB85
C. wasmanni Emery 1893Z01 (see below for synonymy)
C. whitei Wheeler 1915 [=C. (Myrmosphincta) whitei; incl. C. (Myrmosaulus) scutellus Clark 1930]TB85
C. wiederkehri Forel 1894TB85
|--C. w. wiederkehriTB85
`--C. w. lucidior Forel 1910TB85
C. wroughtoni Forel 1893B03
C. yerburyi Forel 1893 [=C. reticulatus yerburyi]B03
C. (Camponotus)T99
C. (Myrmamblys)YT99
C. (Myrmaphaenus) ulcerosus [incl. C. bruesi]WW90
C. (Myrmentoma)YT99
|--C. (M.) caryaeYT99
| |--C. c. caryaeT99
| `--C. c. brunniT99
|--C. (M.) decipiensWBF16
|--C. (M.) keihitoiT99
|--C. (M.) nipponensisT99
|--C. (M.) quadrinotatusT99
`--C. (M.) sayi [incl. C. rasilis]WW90
C. (Myrmocamelus Forel 1914)E14, TB85
|--C. (*M.) ephippium (Smith 1858) [=Formica ephippium]TB85
| |--C. e. ephippiumTB85
| `--C. e. narses Forel 1915TB85
|--C. (M.) gambeyi Emery 1883E14
| |--C. g. gambeyiE14
| `--C. g. marthae For. 1894E14
`--C. (M.) hoplites Emery 1914E14
C. (Myrmosaulus) camelinus (Smith 1857)N-ZLI11, B03 (see below for synonymy)
C. (Myrmotarsus)N-ZLI11
|--C. (M.) irritabilisN-ZLI11
`--C. (M.) rupifemurN-ZLI11
C. (Myrmoturba)E14
|--C. (M.) croceomaculatus Emery 1914E14
|--C. (M.) maculatus (Fabricius 1781)E14, TB85 [=Formica maculataTB85, C. rubripes maculatusG89]
| |--C. m. maculatusTB85
| `--C. m. humilior Forel 1902TB85
`--C. (M.) novaehollandiae Mayr 1870E14, FT08 [=C. maculatus novaehollandiaeE14]
C. (Paramyrmamblys)T99
|--C. (P.) amamianusT99
`--C. (P.) kiusiuensisT99
C. (Tanaemyrmex)YT99
Camponotus angusticollis (Jerdon 1851) [=Formica angusticollis; incl. F. ardens Smith 1858, F. callida Smith 1858, F. impetuosa Smith 1858, Camponotus prismaticus Mayr 1862, C. angusticollis var. sanguinolentus]B03
Camponotus marginatus (Latr. 1798)B03 [=Formica marginataB03; incl. F. glabraL02, Camponotus marginatus var. himalayanus Forel 1893B03]
Camponotus mitis (Smith 1858)B03 [=Formica mitisB03, C. maculatus mitisB03, C. rubripes mitisE89, C. (Myrmoturba) mitisZ01; incl. Formica bacchus Smith 1858B03, C. mitis var. bacchusB03, C. mitis var. comottoi Emery 1887B03, C. mitis var. crassinodis Forel 1892B03, C. mitis var. dulcis Emery 1889B03, C. variegatus var. dulcisT99, C. mitis var. fuscithorax Forel 1892B03, F. ventralis Smith 1858B03]
Camponotus nicobarensis Mayr 1865 [incl. C. nicobarensis exiguoguttatus Forel 1892, C. nicobarensis var. monticola Emery 1894]B03
Camponotus sericeus (Fabr. 1798) [=Formica sericea; incl. F. cinerascens Jerd. 1851, F. obtusa Smith 1858, Camponotus opaciventris Mayr 1878]B03
Camponotus wasmanni Emery 1893Z01 [=C. (Myrmepomis) wasmanniZ01, C. (Orthonotomyrmex) wasmanniZ01; incl. C. wasmanni var. mutilloides Emery 1893B03]
Camponotus (Myrmosaulus) camelinus (Smith 1857)N-ZLI11, B03 [=Formica camelinaB03; incl. F. singularis Smith 1858B03, Camponotus (Myrmosaulus) singularisN-ZLI11]
*Type species of generic name indicated
References
[A61] Andreae, H. 1961. Coleoptera: Cossyphodidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 198–216. Almqvist & Wiksell: Uppsala.
[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera vol. 2. Ants and Cuckoo-Wasps. Taylor and Francis: London.
[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).
[B10] Brown, B. V. 2010. Phoridae (hump-backed flies, scuttle flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 725–761. NRC Research Press: Ottawa.
[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).
[E89] Emery, C. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XX.—Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885–87). Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 485–520.
[E14] Emery, C. 1914. Les fourmis de la Nouvelle-Calédonie & des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 393–436, pl. 12. C. W. Kreidels Verlag: Wiesbaden.
[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.
[FT08] Framenau, V. W., & M. L. Thomas. 2008. Ants (Hymenoptera: Formicidae) of Christmas Island (Indian Ocean): identification and distribution. Records of the Western Australian Museum 25 (1): 45–85.
[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.
[H09] Heterick, B. E. 2009. A guide to the ants of south-western Australia. Records of the Western Australian Museum Supplement 76: 1–206.
[K91] Kathirithamby, J. 1991. Strepsiptera. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 684–695. Melbourne University Press: Carlton (Victoria).
[L02] Latreille, P. A. 1802. Histoire Naturelle des Fourmis, et recueil de mémoires et d’observations sur les abeilles, les araignées, les faucheurs, et autres insectes. Théophile Barrois père: Paris.
[MC05] Machado, G., P. C. Carrera, A. M. Pomini & A. J. Marsaioli. 2005. Chemical defense in harvestmen (Arachnida, Opiliones): do benzoquinone secretions deter invertebrate and vertebrate predators? Journal of Chemical Ecology 31 (11): 2519–2539.
[MS01] Mocsáry, A., & V. Szépligeti. 1901. Hymenopterák [Hymenopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 121–169. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.
[N-ZLI11] Nur-Zati A. M., G. T. Lim & A. B. Idris. 2011. Checklist of ants of selected hill dipterocarp forests of peninsular Malaysia. Serangga 16 (1): 91–105.
[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.
[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.
[SW07] Schöller, M., & V. Witte. 2007. A review of the genus Clytrasoma Jacoby 1908, with description of a new species collected within a Camponotus sp. ant-nest (Insecta: Coleoptera, Chrysomelidae; Hymenoptera, Formicidae). Senckenbergiana Biologica 87 (1): 51–61.
[TB85] Taylor, R. W., & D. R. Brown. 1985. Formicoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 1–149. Australian Government Publishing Service: Canberra.
[T99] Terayama, M. 1999. The ant genus Camponotus Mayr (Hymenoptera: Formicidae) in Japan. Memoirs of the Myrmecological Society of Japan 1: 25–48.
[WL04] Wang, X.-H., & P. J. Lester. 2004. A preliminary study of the usefulness of morphometric tools for splitting the Monomorium antarcticum (Smith) complex (Hymenoptera: Formicidae), New Zealand’s most common native ants. New Zealand Entomologist 27: 103–108.
[WBF16] Ward, P. S., B. B. Blaimer & B. L. Fisher. 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.
[WB03] Ward, P. S., & S. G. Brady. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.
Wernegreen, J. J., S. N. Kauppinen, S. G. Brady & P. S. Ward. 2009. One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology 9: 292. doi:10.1186/1471-2148-9-292.
[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.
[YT99] Yamane, S., & M. Terayama. 1999. A new species of the genus Pristomyrmex Mayr from Japan, and a proposal of a new synonym of species in the genus Camponotus Mayr (Hymenoptera: Formicidae). Memoirs of the Myrmecological Society of Japan 1: 17–24.
[Z01] Zhou, S. 2001. Ants of Guangxi. Guangxi Normal University Press: Guilin (China).