Camponotus quadrimaculatus

AntWiki: The Ants --- Online
Camponotus quadrimaculatus
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Tribe: Camponotini
Genus: Camponotus
Subgenus: Myrmosaga
Species: C. quadrimaculatus
Binomial name
Camponotus quadrimaculatus
Forel, 1886

Camponotus quadrimaculatus casent0146549 profile 1.jpg

Camponotus quadrimaculatus casent0146549 dorsal 1.jpg

Specimen labels

Synonyms

Endemic to the Malagasy region, C. quadrimaculatus is known from Madagascar, the Comoros islands (Anjouan, Grand Comore and Moheli), and Mayotte. In Madagascar, this species is widely distributed across most of the terrestrial habitats of the island except spiny bush and thicket in the southern region. Camponotus quadrimaculatus is typically found from littoral rainforest and dry forest through rainforest and montane rainforests to savannah woodland habitats. It also can colonize human-modified habitats adjacent to its natural areas of distribution. The yellowish orange variant of the species has a range restricted to the northern part of Madagascar, where it occupies not only natural rainforest areas of the RS Manongarivo and the Galoko Montain Chains and the littoral rainforest of Nosy Faly, but also human-modified areas such as coffee plantations, urban garden habitats, and disturbed littoral rainforest habitats. (Rakotonirina & Fisher, 2022)

Identification

Rakotonirina and Fisher (2022) - In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; two pairs of white spots present on second and third abdominal tergites; pronotum, mesonotum, and propodeum not forming separate convexities, metanotal groove not depressed; propodeum immediately in junction with promesonotum; propodeal dorsum concave.

Camponotus quadrimaculatus is similar to Camponotus rotrae but in the latter the propodeal dorsum is straight.

Keys including this Species

Distribution

Latitudinal Distribution Pattern

Latitudinal Range: -11.47435° to -24.95694°.

 
North
Temperate 		North
Subtropical 		Tropical South
Subtropical 		South
Temperate

Distribution based on Regional Taxon Lists

Afrotropical Region: Comoros.
Malagasy Region: Madagascar (type locality), Mayotte.

Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
pChart

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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Biology

Castes

Worker

Images from AntWeb

Camponotus quadrimaculatus casent0102426 head 1.jpgCamponotus quadrimaculatus casent0102426 profile 1.jpgCamponotus quadrimaculatus casent0102426 dorsal 1.jpgCamponotus quadrimaculatus casent0102426 label 1.jpg
Lectotype of Camponotus quadrimaculatus. Worker. Specimen code casent0102426. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus casent0102427 head 1.jpgCamponotus quadrimaculatus casent0102427 profile 1.jpgCamponotus quadrimaculatus casent0102427 dorsal 1.jpgCamponotus quadrimaculatus casent0102427 label 1.jpg
Paralectotype of Camponotus quadrimaculatus. Worker (major/soldier). Specimen code casent0102427. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus casent0102442 head 1.jpgCamponotus quadrimaculatus casent0102442 profile 1.jpgCamponotus quadrimaculatus casent0102442 dorsal 1.jpgCamponotus quadrimaculatus casent0102442 label 1.jpg
Paralectotype of Camponotus quadrimaculatus. Worker (major/soldier). Specimen code casent0102442. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus casent0102443 head 1.jpgCamponotus quadrimaculatus casent0102443 profile 1.jpgCamponotus quadrimaculatus casent0102443 dorsal 1.jpgCamponotus quadrimaculatus casent0102443 label 1.jpg
Paralectotype of Camponotus quadrimaculatus. Worker. Specimen code casent0102443. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus casent0133564 head 1.jpgCamponotus quadrimaculatus casent0133564 profile 1.jpgCamponotus quadrimaculatus casent0133564 dorsal 1.jpgCamponotus quadrimaculatus casent0133564 label 1.jpg
Worker (major/soldier). Specimen code casent0133564. Photographer Erin Prado, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Camponotus quadrimaculatus casent0134917 head 1.jpgCamponotus quadrimaculatus casent0134917 profile 1.jpgCamponotus quadrimaculatus casent0134917 dorsal 1.jpgCamponotus quadrimaculatus casent0134917 label 1.jpg
Worker. Specimen code casent0134917. Photographer Erin Prado, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Camponotus quadrimaculatus casent0145975 head 1.jpgCamponotus quadrimaculatus casent0145975 profile 1.jpgCamponotus quadrimaculatus casent0145975 dorsal 1.jpgCamponotus quadrimaculatus casent0145975 label 1.jpg
Worker (major/soldier). Specimen code casent0145975. Photographer Erin Prado, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Camponotus kelleri casent0101519 head 1.jpgCamponotus kelleri casent0101519 profile 1.jpgCamponotus kelleri casent0101519 dorsal 1.jpgCamponotus kelleri casent0101519 label 1.jpg
Lectotype of Camponotus kelleri. Worker (major/soldier). Specimen code casent0101519. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus kelleri casent0101524 head 1.jpgCamponotus kelleri casent0101524 profile 1.jpgCamponotus kelleri casent0101524 dorsal 1.jpgCamponotus kelleri casent0101524 label 1.jpg
Paralectotype of Camponotus kelleri. Worker (major/soldier). Specimen code casent0101524. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus kelleri casent0101525 head 1.jpgCamponotus kelleri casent0101525 profile 1.jpgCamponotus kelleri casent0101525 dorsal 1.jpgCamponotus kelleri casent0101525 label 1.jpg
Paralectotype of Camponotus kelleri. Worker (major/soldier). Specimen code casent0101525. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus kelleri invalidus casent0101517 head 1.jpgCamponotus kelleri invalidus casent0101517 profile 1.jpgCamponotus kelleri invalidus casent0101517 dorsal 1.jpgCamponotus kelleri invalidus casent0101517 label 1.jpg
Lectotype of Camponotus kelleri invalidus. Worker. Specimen code casent0101517. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus kelleri invalidus casent0101520 head 1.jpgCamponotus kelleri invalidus casent0101520 profile 1.jpgCamponotus kelleri invalidus casent0101520 dorsal 1.jpgCamponotus kelleri invalidus casent0101520 label 1.jpg
Paralectotype of Camponotus kelleri invalidus. Worker (major/soldier). Specimen code casent0101520. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus sellaris casent0102109 head 1.jpgCamponotus quadrimaculatus sellaris casent0102109 profile 1.jpgCamponotus quadrimaculatus sellaris casent0102109 dorsal 1.jpgCamponotus quadrimaculatus sellaris casent0102109 label 1.jpg
Lectotype of Camponotus quadrimaculatus sellaris. Worker. Specimen code casent0102109. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MSNG, Genoa, Italy.
Camponotus quadrimaculatus sellaris casent0102110 head 1.jpgCamponotus quadrimaculatus sellaris casent0102110 profile 1.jpgCamponotus quadrimaculatus sellaris casent0102110 profile 2.jpgCamponotus quadrimaculatus sellaris casent0102110 dorsal 1.jpgCamponotus quadrimaculatus sellaris casent0102110 label 1.jpg
Paralectotype of Camponotus quadrimaculatus sellaris. Worker (major/soldier). Specimen code casent0102110. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MSNG, Genoa, Italy.
Camponotus quadrimaculatus sellaris casent0102111 head 1.jpgCamponotus quadrimaculatus sellaris casent0102111 profile 1.jpgCamponotus quadrimaculatus sellaris casent0102111 dorsal 1.jpgCamponotus quadrimaculatus sellaris casent0102111 label 1.jpg
Paralectotype of Camponotus quadrimaculatus sellaris. Worker (major/soldier). Specimen code casent0102111. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MSNG, Genoa, Italy.
Camponotus quadrimaculatus sellaris casent0102435 head 1.jpgCamponotus quadrimaculatus sellaris casent0102435 profile 1.jpgCamponotus quadrimaculatus sellaris casent0102435 profile 2.jpgCamponotus quadrimaculatus sellaris casent0102435 dorsal 1.jpgCamponotus quadrimaculatus sellaris casent0102435 label 1.jpg
Paralectotype of Camponotus quadrimaculatus sellaris. Worker (major/soldier). Specimen code casent0102435. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus sellaris casent0102436 head 1.jpgCamponotus quadrimaculatus sellaris casent0102436 profile 1.jpgCamponotus quadrimaculatus sellaris casent0102436 dorsal 1.jpgCamponotus quadrimaculatus sellaris casent0102436 label 1.jpg
Paralectotype of Camponotus quadrimaculatus sellaris. Worker. Specimen code casent0102436. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.

Queen

Images from AntWeb

Camponotus quadrimaculatus casent0102424 head 1.jpgCamponotus quadrimaculatus casent0102424 profile 1.jpgCamponotus quadrimaculatus casent0102424 dorsal 1.jpgCamponotus quadrimaculatus casent0102424 label 1.jpg
Queen (alate/dealate). Specimen code casent0102424. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by Museum of Natural History, Geneva.
Camponotus quadrimaculatus casent0133512 head 1.jpgCamponotus quadrimaculatus casent0133512 profile 1.jpgCamponotus quadrimaculatus casent0133512 dorsal 1.jpgCamponotus quadrimaculatus casent0133512 label 1.jpg
Queen (alate/dealate). Specimen code casent0133512. Photographer Erin Prado, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.
Camponotus quadrimaculatus casent0147466 head 1.jpgCamponotus quadrimaculatus casent0147466 profile 1.jpgCamponotus quadrimaculatus casent0147466 profile 2.jpgCamponotus quadrimaculatus casent0147466 dorsal 1.jpgCamponotus quadrimaculatus casent0147466 label 1.jpg
Queen (alate/dealate). Specimen code casent0147466. Photographer Erin Prado, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.

Male

Images from AntWeb

Camponotus quadrimaculatus casent0102425 head 1.jpgCamponotus quadrimaculatus casent0102425 profile 1.jpgCamponotus quadrimaculatus casent0102425 dorsal 1.jpgCamponotus quadrimaculatus casent0102425 label 1.jpg
Paralectotype of Camponotus quadrimaculatus. Male (alate). Specimen code casent0102425. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by MHNG, Geneva, Switzerland.
Camponotus quadrimaculatus casent0104625 profile 1.jpgCamponotus quadrimaculatus casent0104625 dorsal 1.jpgCamponotus quadrimaculatus casent0104625 label 1.jpg
Paralectotype of Camponotus quadrimaculatus. Male (alate). Specimen code casent0104625. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ZMHB, Berlin, Germany.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • quadrimaculatus. Camponotus quadrimaculatus Forel, 1886c: cii (s.w.) MADAGASCAR.
    • Type-material: syntype major and minor workers (numbers not stated).
    • Type-locality: Madagascar: (no further data) (A. Grandidier).
    • Type-depository: probably MHNG.
    • [Notes (i): the designation by Rakotonirina & Fisher, 2022: 140, consisting of lectotype minor worker, 2 paralectotype major workers, 1 paralectotype minor worker, 1 paralectotype queen, 2 paralectotype males, Madagascar: Fianarantsoa (L. Besson), is redundant as this locality and collector do not appear in the original description; (ii) the Besson specimens apparently derive from Forel, 1891b: 59.]
    • [Camponotus quadrimaculatus Forel, 1879a: 115. Nomen nudum.]
    • [Misspelled as 4 maculatus by Forel, 1907g: 91.]
    • Forel, 1891b: 59 (q.m.).
    • Combination in C. (Mayria): Emery, 1925b: 123;
    • combination in C. (Myrmosaga): Forel, 1912i: 92; Rakotonirina & Fisher, 2022: 140.
    • Status as species: Forel, 1891b: 58 (redescription); Dalla Torre, 1893: 249; Emery, 1896d: 374 (in list); Forel, 1907g: 91; Wheeler, W.M. 1922a: 1046; Emery, 1925b: 123; Bolton, 1995b: 119; Rakotonirina & Fisher, 2022: 140 (redescription).
    • Senior synonym of invalidus: Rakotonirina & Fisher, 2022: 141.
    • Senior synonym of kelleri: Rakotonirina & Fisher, 2022: 141.
    • Senior synonym of sellaris: Rakotonirina & Fisher, 2022: 140.
    • Distribution: Comoros, Madagascar, Mayotte.
  • invalidus. Camponotus kelleri var. invalidus Forel, 1897c: 200 (s.w.) MADAGASCAR.
    • Type-material: lectotype minor worker (by designation of Rakotonirina & Fisher, 2022: 141), 1 paralectotype major worker.
    • Type-locality: lectotype Madagascar: Nossi-Bé (A. Voeltzkow); paralectotype with same data.
    • Type-depository: MHNG.
    • Combination in C. (Mayria): Emery, 1925b: 123.
    • Subspecies of kelleri: Emery, 1898a: 226; Wheeler, W.M. 1922a: 1046; Emery, 1925b: 123; Bolton, 1995b: 105.
    • Junior synonym of quadrimaculatus: Rakotonirina & Fisher, 2022: 141.
  • kelleri. Camponotus kelleri Forel, 1886f: 186 (s.w.) MADAGASCAR.
    • Type-material: lectotype major worker (by designation of Rakotonirina & Fisher, 2022: 141), 2 paralectotype major workers.
    • Type-locality: lectotype Madagascar: nr Tamatavé (C. Keller); paralectotypes with same data.
    • Type-depository: MHNG.
    • Combination in C. (Myrmosaga): Forel, 1912i: 92;
    • combination in C. (Mayria): Emery, 1925b: 123.
    • Status as species: Forel, 1891b: 62 (redescription); Dalla Torre, 1893: 237; Emery, 1895f: 337; Emery, 1896d: 374 (in list); Emery, 1898a: 226; Forel, 1907e: 14; Wheeler, W.M. 1922a: 1046; Emery, 1925b: 123; Bolton, 1995b: 106.
    • Junior synonym of quadrimaculatus: Rakotonirina & Fisher, 2022: 141.
  • sellaris. Camponotus quadrimaculatus subsp. sellaris Emery, 1895f: 344, fig. 3 (w.) MADAGASCAR.
    • Type-material: lectotype minor worker (by designation of Rakotonirina & Fisher, 2022: 140), 3 paralectotype major workers, 1 paralectotype minor worker.
    • Type-locality: lectotype Madagascar: Diego Suarez (= Antsiranana), iv.-viii.1893 (Ch. Alluaud); paralectotypes with same data.
    • Type-depositories: MSNG (lectotype); MHNG, MSNG (paralectotypes).
    • Combination in C. (Mayria): Emery, 1925b: 123.
    • Subspecies of quadrimaculatus: Emery, 1896d: 374 (in list); Wheeler, W.M. 1922a: 1047; Emery, 1925b: 123; Bolton, 1995b: 123.
    • Junior synonym of quadrimaculatus: Rakotonirina & Fisher, 2022: 140.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

Rakotonirina and Fisher (2022):

Morphological measurements: see Appendix 1 and Ratios of morphometric data for majors and minors

Minor In full-face view, head widest at midlength, lateral margins posterior to level of eye rounding evenly to posterior margin; eye large, slightly convex (EL/CS: 0.27±0.01; 0.26–0.30), not breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/4 of the head (PoOc/CL: 0.28±0.02; 0.25–0.32); frontal carinae wide (FR/CS: 0.25±0.01; 0.24–0.28), distance between them larger than their distance to eye; clypeus with anterolateral angle and anteromedian margin with blunt or convex angle; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.70±0.11; 1.59–1.88). Promesonotum weakly convex, posterior portion of mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum almost straight; dorsal margin of propodeum and declivity junction bluntly rounded; propodeal dorsum 2 × as long as declivity. Petiolar node short and high, its dorsal margin inclined posteriorly and joining anterior face in a blunt angle; anterior face 1/3 of height of the posterior; femur of hind leg rounded axially, without torsion near base.

First and second gastral tergites without a pair of white spots; erect hairs present on lateral margin of head, near its posterior margin bearing more than six erect hairs; antennal scape with suberect hairs inclined at ca. 30°, appressed hairs lacking; promesonotum covered with erect hairs; posterodorsal angle of propodeum with more than two erect hairs.

Major Differing from minor worker in the following characters: enlarged head (CS: 4.02±0.35; 3.69–4.61; CWb/CL: 0.92±0.08; 0.79–1.01); more robust mandible; apical 1/4 of antennal scape surpassing posterior cephalic margin; pronotum convex, mesonotum sloping towards metanotum, propodeal dorsum feebly convex and joining the declivity at a broad angle; petiolar node higher than long.

Type Material

Camponotus kelleri

  • Syntype workers, Madagascar, Toamasina (C. Keller); 1 syntype major worker designated as lectotype, by present designation, AntWeb CASENT0101519 (MHNG) [Rakotonirina & Fisher, 2022].
  • Paralectotypes: 2 major workers of same data as lectotype but with the following specimen codes: CASENT0101524, CASENT0101525 (MHNG) [Rakotonirina & Fisher, 2022].

Camponotus kelleri invalidus

  • Syntype workers, Madagascar, Nosibé [Antsiranana, Nosy be, -13.291667, 48.258335, 146 m] (Voeltzkow); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101517 (MHNG) [Rakotonirina & Fisher, 2022].
  • Paralectotype: 1 major worker of same data as lectotype but specimen coded as: CASENT0101520 (MHNG) [Rakotonirina & Fisher, 2022].

Camponotus quadrimaculatus

  • Lectotype minor worker, by designation of Rakotonirina & Fisher (2022), Madagascar, Fianarantsoa (Besson) AntWeb CASENT0102426 (MHNG) [Rakotonirina & Fisher, 2022].
  • Paralectotypes: 1 minor worker, 2 major workers, 1 alate queen and 2 males of same data as lectotype but respectively specimen coded as: CASENT0102443, CASENT0102427, CASENT0102442 CASENT0102424 (queen), CASENT0102425 (MHNG), CASENT0104625 (ZMHB) (male) (Besson).

Camponotus quadrimaculatus sellaris

  • Syntype workers, Madagascar, Antsiranana [[Diego Suarez; 7]; Antsahampano; Antsiranana Rural, -12.323135, 49.294285, 67 m] (Alluaud 1893); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102109 (MSNG) [Rakotonirina & Fisher, 2022].
  • Paralectotypes: 1 minor worker, 3 major workers of same data as lectotype but respectively with the following specimen codes: CASENT0102436 (MHNG); CASENT0102110, CASENT0102111 (MSNG); CASENT0102435 (MHNG) [examined]. [Combination in Camponotus (Mayria) Emery, 1925: 123]. Syn. nov.

Taxonomic Notes

The minor workers of C. quadrimaculatus display a remarkable range of phenotypic diversity. This assertion of the diversity of morphological characters is made on the basis of specimens with similar promesonotum shape, dorsal outline of the propodeal dorsum, and color of the body. However, the geographic range of differences among the observed variants does not follow a simple pattern. One variant of the yellowish orange body color has a restricted geographic boundary in the north of Madagascar, while others, showing variation in the strength of the concavity on the propodeal dorsum, are widely distributed along the rainforests of the island.

Across the wide geographic range of Camponotus quadrimaculatus, trait variations in the worker castes of C. quadrimaculatus sellaris, Camponotus kelleri and Camponotus kelleri invalidus have been observed. However, these variations do not present any pattern that separates them from the former species. Accordingly, C. quadrimaculatus sellaris, Camponotus kelleri and Camponotus kelleri invalidus are synonymyzed under Camponotus quadrimaculatus (Rakotonirina & Fisher, 2022).

References

  • Emery, C. 1925d. Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum 183: 1-302 (page 123, Combination in C. (Mayria))
  • Forel, A. 1886c. Diagnoses provisoires de quelques espèces nouvelles de fourmis de Madagascar, récoltées par M. Grandidier. Ann. Soc. Entomol. Belg. 30:ci-cvii. (page cii, soldier, worker described)
  • Forel, A. 1891c. Les Formicides. [part]. In: Grandidier, A. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (28e fascicule). Paris: Hachette et Cie, v + 237 pp. (page 58, soldier, queen, male described)
  • Forel, A. 1912j. Formicides néotropiques. Part VI. 5me sous-famille Camponotinae Forel. Mém. Soc. Entomol. Belg. 20: 59-92 (page 92, Combination in C. (Myrmosaga))
  • Rakotonirina, J.C., Fisher, B.L. 2022. Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology. ZooKeys 1098: 1–180 (doi:10.3897/zookeys.1098.73223).
  • Wheeler, W. M. 1922k. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. IX. A synonymic list of the ants of the Malagasy region. Bull. Am. Mus. Nat. Hist. 4 45: 1005-1055 (see also)

References based on Global Ant Biodiversity Informatics

  • Emery C. 1895. Mission scientifique de M. Ch. Alluaud dans le territoire de Diego-Suarez (Madagascar-nord) (Avril-août 1893). Formicides. Annales de la Société Entomologique de Belgique 39: 336-345.
  • Fisher B. L. 1997. Biogeography and ecology of the ant fauna of Madagascar (Hymenoptera: Formicidae). Journal of Natural History 31: 269-302.
  • Fisher B. L. 2003. Formicidae, ants. Pp. 811-819 in: Goodman, S. M.; Benstead, J. P. (eds.) 2003. The natural history of Madagascar. Chicago: University of Chicago Press, xxi + 1709 pp.
  • Forel A. 1886. Diagnoses provisoires de quelques espèces nouvelles de fourmis de Madagascar, récoltées par M. Grandidier. Annales de la Société Entomologique de Belgique. 30: ci-cvii.
  • Forel A. 1887. Fourmis récoltées à Madagascar par le Dr. Conrad Keller. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 7: 381-389.
  • Forel A. 1897. Ameisen aus Nossi-Bé, Majunga, Juan de Nova (Madagaskar), den Aldabra-Inseln und Sansibar, gesammelt von Herrn Dr. A. Voeltzkow aus Berlin. Mit einem Anhang über die von Herrn Privatdocenten Dr. A. Brauer in Marburg auf den Seychellen und von Herrn Perrot auf Ste. Marie (Madagaskar) gesammelten Ameisen. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 21: 185-208.
  • Forel A. 1907. Ameisen von Madagaskar, den Comoren und Ostafrika. Wissenschaftliche Ergebnisse. Reise in Ostafrika 2: 75-92.
  • Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. IX. A synonymic list of the ants of the Malagasy region. Bulletin of the American Museum of Natural History 45: 1005-1055
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