Camponotus quadrimaculatus

Endemic to the Malagasy region, C. quadrimaculatus is known from Madagascar, the Comoros islands (Anjouan, Grand Comore and Moheli), and Mayotte (Fig. 69D). 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
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.

Distribution based on Regional Taxon Lists
Afrotropical Region: Comoros. Malagasy Region: Madagascar, Mayotte.

Nomenclature

 * . Camponotus quadrimaculatus Forel, 1886c: cii (s.w.) MADAGASCAR.
 * [Camponotus quadrimaculatus Forel, 1879a: 115. Nomen nudum.]
 * [Misspelled as 4 maculatus by Forel, 1907g: 91.]
 * Forel, 1891b: 59 (q.m.).
 * Combination in C. (Myrmosaga): Forel, 1912i: 92;
 * combination in C. (Mayria): Emery, 1925b: 123.
 * 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.
 * kelleri. Camponotus kelleri Forel, 1886f: 186 (s.w.) MADAGASCAR.
 * 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.
 * invalidus. Camponotus kelleri var. invalidus Forel, 1897c: 200 (w.) MADAGASCAR.
 * 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.
 * sellaris. Camponotus quadrimaculatus subsp. sellaris Emery, 1895f: 344, fig. 3 (w.) MADAGASCAR.
 * 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.

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 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