Camponotus quadrimaculatus Forel, 1886

Rakotonirina, Jean Claude & Fisher, Brian L., 2022, Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology, ZooKeys 1098, pp. 1-180 : 1

publication ID	https://dx.doi.org/10.3897/zookeys.1098.73223
publication LSID	lsid:zoobank.org:pub:B4F4033F-296E-43CC-BE54-B9413BC19268
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scientific name	Camponotus quadrimaculatus Forel
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Camponotus quadrimaculatus Forel

Figs 15C View Figure 15 , 24B View Figure 24 , 35B View Figure 35 , 38B View Figure 38 , 39B View Figure 39 , 69 View Figure 69

Camponotus quadrimaculatus Forel, 1886a: cii. Lectotype minor worker, by present designation, Madagascar, Fianarantsoa (Besson) AntWeb CASENT0102426 ( MHNG) [examined]. 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, CASENT0102442CASENT0102424 (queen), CASENT0102425 ( MHNG), CASENT0104625 (ZMHB) (male) (Besson). [ Camponotus quadrimaculatus Forel, 1879: 115. Nomen nudum.]. Raised to species by Forel 1891: 58 (redescription), 1907: 91; Dalla Torre 1893: 249; Emery 1896: 374, 1925: 123; Wheeler 1922: 1046; Bolton 1995: 119. [Combination in Camponotus (Myrmosaga) Forel, 1912: 92; in Camponotus (Mayria) Emery, 1925: 123].

Camponotus quadrimaculatus sellaris Emery, 1895: 344. 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) [examined]. 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.

Camponotus kelleri Forel, 1886b: 186. Syntype workers, Madagascar, Toamasina (C. Keller); 1 syntype major worker designated as lectotype, by present designation, AntWeb CASENT0101519 ( MHNG) [examined]. Paralectotypes: 2 major workers of same data as lectotype but with the following specimen codes: CASENT0101524, CASENT0101525 ( MHNG) [examined]. Syn. nov.

Camponotus kelleri var. invalidus Forel, 1897: 200. 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) [examined]. Paralectotype: 1 major worker of same data as lectotype but specimen coded as: CASENT0101520 ( MHNG) [examined]. [Combination in Camponotus (Mayria) : Emery, 1925: 123]. Syn. nov.

Additional material examined.

Comoros: Anjouan: Lac Dzialandée, -12.22474, 44.43121, 900 m, disturbed montane rainforest (B.L. Fisher et al.) ( CAS). Mount Ntringui, -12.22043, 44.42924, 1225 m, montane forest (B.L. Fisher et al.) ( CAS). -12.18771, 44.35929, 65 m, coastal roadside (B.L. Fisher et al.) ( CAS). -12.30537, 44.45031, 500 m, along roadside, mango, banana (B.L. Fisher et al.) ( CAS). -12.30355, 44.46926, 885 m, along roadside plantation cows (B.L. Fisher et al.) ( CAS). -12.29311, 44.5109, 440 m, along roadside, mango (B.L. Fisher et al.) ( CAS). Grande Comore: Grillé, -11.47578, 43.34669, 995 m, montane rainforest (B.L. Fisher et al.) ( CAS); Grillé, -11.47578, 43.34669, 805 m, bananas plantation (B.L. Fisher et al.) ( CAS). Itoundzou, -11.63136, 43.30434, 635 m, secondary rainforest along roadside (B.L. Fisher et al.) ( CAS). Karthala, -11.82699, 43.4295, 1000 m, montane rainforest (B.L. Fisher et al.) ( CAS). Mouadja, -11.47435, 43.3004, 350 m, coastal scrub (B.L. Fisher et al.) ( CAS). Mohéli: Ouallah, -12.32717, 43.65952, 10 m, coastal scrub (B.L. Fisher et al.) ( CAS); Ouallah, -12.30668, 43.66407, 275 m, rainforest (B.L. Fisher et al.) ( CAS); Ouallah, -12.29696, 43.67392, 680 m, rainforest (B.L. Fisher et al.) ( CAS). Madagascar: Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle (Mike, Rinha) ( CAS); Ankokoy Forest, 3 km E Ibity, malaise in Uapaca forest, -20.0675, 46.9995, 1700 m, Uapaca forest (M.E. Irwin, Rin’ha) ( CAS). Antsiranana: [ Nosibé], Nosy be, Nosibe, -13.29166667, 48.25833333, 146 m ( MHNG); [Diego Suarez; 7]; Antsahampano; Antsiranana Rural, -12.323135, 49.294283, 67 m (Ch. Alluaud) ( MSNG); Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) ( CAS); Ambondrobe, 41.1 km 175° NW Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest (B.L. Fisher) ( CAS); Forêt Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) ( CAS); Forêt d’Antsahabe, 11.4 km 275° W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher) ( CAS); Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) ( CAS); Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 650-800 m, rainforest (B.L. Fisher) ( CAS); Galoko chain, Mont Galoko, -13.58487, 48.71818, 520 m, rainforest (B.L. Fisher et al.) ( CAS); PN Montagne d’Ambre, Pic Bades, -12.5186, 49.18625, 900 m, montane rainforest (B.L. Fisher et al.) ( CAS); Antongombato, 2.2 km SW Antsiranana, -12.37277, 49.22888, 74 m, urban/garden (B.L. Fisher et al.) ( CAS); RS Manongarivo, 14.5 km 220° SW Antanambao, -13.99833, 48.42833, 1175 m, montane rainforest (B.L. Fisher) ( CAS); Réserve Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) ( CAS); RS Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); Sakaramy 07 km N of Joffre Ville, -12.33333, 49.25, 360 m, Low rain forest in open area (Mike, Frank, Rin’ha) ( CAS). Fianarantsoa: [Fianarantsoa], 2eme Arrondissement; Fianarantsoa Urban, -21.43333, 47.08333, 1160 m (Dr Besson) ( MHNG); Amoron’i Mania Region, District of Ambositra, Italaviana Uapaca forest, 35 km SE of Antsirabe, -20.17333, 47.086, 1359 m, Uapaca forest ( Rin’Ha, Mike) ( CAS); Ampangabe V Non Protected Area, 21.37 km W Itremo, -20.61361, 46.60799, 1449 m, Shrubland (A. Ravelomanana) ( CAS); Ampangabe VI Non Protected Area, 21.16 km W Itremo, -20.61444, 46.6104, 1379 m, Shrubland (A. Ravelomanana) ( CAS); dry wash, 1 km E of PN Isalo Interpretive Center, -22.62667, 45.35817, 885 m, dry wash (M.E. Irwin, F.D. Parker, R. Harin’Hala) ( CAS); Forêt d’Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); Horombe Region, District of Ihosy, Betapia (Border of Fianarantsoa and Tulear): 09 km SW of Ilakaka Saphir town, -22.62883, 45.36117, 1036 m, Uapaca forest ( Rin’Ha, Mike) ( CAS); RS Manombo, 32 km SE of Farafangana, -23.02183, 47.72, 36 m, Lowland rainforest ( Rin’Ha, Mike) ( CAS); PN Isalo, 9.1 km 354° N Ranohira, -22.48167, 45.46167, 725 m, gallery forest (Fisher, Griswold et al.) ( CAS); PN Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland (Fisher, Griswold et al.) ( CAS); PN Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) ( CAS); stream area, 900 m E of PN Isalo Interpretive Center, -22.62667, 45.35817, 750 m, open area near stream (R. Harin’Hala) ( CAS); Tsaranoro, 32.8 km 229° Ambalavao, -22.08483, 46.77633, 950 m, rainforest (B.L. Fisher et al.) ( CAS); Tsaranoro, 32.8 km 230° Ambalavao, -22.08317, 46.774, 975 m, savannah woodland (B.L. Fisher et al.) ( CAS); Vohiparara broken bridge, -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin’Hala) ( CAS). Mahajanga: Boeny Region, District of Marovoay, PN Ankarafantsika, Ampijoroa SF, 160 km North of Maevatanana on RN 04, -16.31933, 46.81333, 42 m, deciduous forest ( Rin’Ha, Mike) ( CAS); Region Sofia, Bemanevika, -14.33886, 48.58729, 1567 m, montane rainforest (B.L. Fisher et al.) ( CAS); Region Sofia, Bemanevika, -14.337, 48.58874, 1606 m, montane rainforest (B.L. Fisher et al.) ( CAS); RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transition humid forest (B.L. Fisher et al.) ( CAS); Sofia Region, District of Port-Berger, Ambovomamy 20 km N of Port-Berger, -15.45117, 47.61333, 86 m, secondary forest on white sandy area (Mike, Frank, Rin’ha) ( CAS); PN Ankarafantsika, Ampijoroa SF, 160 km N Maevatanana, deciduous forest, -16.31944, 46.81333, 43 m, deciduous forest (M. Irwin and Rin’ha Harin’hala) ( CAS); Boeny Region, District of Marovoay, PN Ankarafantsika, Ampijoroa SF, 160 km North of Maevatanana on RN 04, -16.31933, 46.81333, 42 m, deciduous forest ( Rin’Ha, Mike) ( CAS); Forêt Ambohimanga, 26.1 km 314° Mampikony, -15.96267, 47.43817, 250 m, tropical dry forest (B.L. Fisher) ( CAS); Mahavavy River, 6.2 km 145° SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) ( CAS); PN Ankarafantsika, Ampijoroa SF, 40 km 306° NW Andranofasika, -16.32083, 46.81067, 130 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); PN Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); PN Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); PN Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) ( CAS); Réserve d’Ankoririka, 10.6 km 13° NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); RS Bemarivo, 23.8 km 223° SW Besalampy, -16.925, 44.36833, 30 m, tropical dry forest (Fisher, Griswold et al.) ( CAS); Sofia Region, District of Port-Berger, Ambovomamy 20 km N of Port-Berger, -15.45117, 47.61333, 86 m, secondary forest on white sandy area (Mike, Frank, Rin’ha) ( CAS); Station Forestiere Ampijoroa, -16.31667, 46.81667, 80 m, tropical dry forest (P.S. Ward) ( CAS); Mampikony, -16.09323, 47.64278, 49 m, urban/garden (B.L. Fisher et al.) ( CAS). Toamasina: 25 km W. Morarano-Chrome, -17.75, 47.98333, 1200 m (A. Pauly) ( CAS); Besarikata, env. Réserve Zahamena, -17.45, 48.85 (A. Pauly) ( CAS). Forêt Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (B.L. Fisher) ( CAS); Nosy Mangabe, -15.5, 49.76667, <5 m, rainforest edge (P.S. Ward) ( CAS). Toliara: Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 8 km NW of Amboasary, Berenty Special Reserve, -25.00667, 46.30333, 85 m, Galery forest ( Rin’Ha, Mike) ( CAS); Atsimo Andrefana Region, District of Horombe, Vohibasia National Parc, Ambakaka Galery forest, 44 km NE of Sakaraha, -23.38183, 43.71267, 182 m, palm forest (Mike, Rinha) ( CAS); Beza-Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) ( CAS); Makay Mts., -21.21985, 45.32396, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) ( CAS); Makay Mts., -21.22336, 45.32628, 480 m, gallery forest on sandy soil (B.L. Fisher et al.) ( CAS); Makay Mts., -21.21761, 45.33917, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) ( CAS); 3 km NE Ambohitra (=Joffreville), -12.46667, 49.21667, 520 m, rainforest edge (P.S. Ward) ( CAS); Makay, -21.31477, 45.12926, 600 m, deciduous dry forest (J.M. Bichain) ( CAS); Makay, -21.3133, 45.14788, 520 m, gallery forest (J.M. Bichain) ( CAS); 1 km E Mahamavo, PN Andohahela, -24.76667, 46.73333, 600 m, rainforest edge (P.S. Ward) ( CAS). Mayotte: Chembenyoumba, -12.762222, 45.075278 (R. Jocqué) ( CAS); Chembenyoumba, Tsanaraki plage, -12.75, 45.06667, litter of shrubs on mangrove edge (R. Jocqué & G. DeSmet) ( CAS); Coconi, DAF Campus, -12.83333, 45.13333 (R. Jocqué) ( CAS); Coconi, DAF Campus, -12.83333, 45.13333 (R. Jocqué & G. DeSmet) ( CAS); Coconi, SDA (service du develppement agricole), -12.83333, 45.13333, wet area with giant bamboo (R. Jocqué & G. DeSmet) ( CAS); Coconi, SDA (service du développement agricole), -12.83333, 45.13333, forested area with giant bamboo (R. Jocqué & G. DeSmet) ( CAS); Dziani Karihani, -12.78333, 45.11667, forest (R. Jocqué & G. DeSmet) ( CAS); Dziani Karihani, Ilang-Ilang plantation, -12.78333, 45.11667 (R. Jocqué & G. DeSmet) ( CAS); entre Combani and Bouyouni, -12.735556, 45.138333, plantation (R. Jocqué) ( CAS); Majimbini, -12.765833, 45.188056, in ruins (R. Jocqué) ( CAS); Majimbini, -12.765833, 45.188056 (R. Jocqué) ( CAS); Majimbini, -12.765833, 45.188056 (R. Jocqué) ( CAS); Mlima Choungi, near crossroad GRMT1 & CCT11 (R. Jocqué) ( CAS); Mlima Choungi, near crossroad GRMT1 & CCT11 (R. Jocqué) ( CAS); Mont Benara, -12.87585, 45.15672, 425 m, rainforest (B.L. Fisher et al.) ( CAS); Mont Chongui, -12.95903, 45.13411, 380 m, rainforest (B.L. Fisher et al.) ( CAS); Mont Chongui summit, -12.99567, 45.13428, 550 m, rainforest (B.L. Fisher et al.) ( CAS); Mont Combani, -12.80632, 45.15314, 370 m, rainforest (B.L. Fisher et al.) ( CAS); Mt. Choungui, -12.8, 45.1, 360 m, forest near fallen tree (R. Jocqué & G. DeSmet) ( CAS); Reserve Forestière Majimbini, -12.76796, 45.18615, 525 m, rainforest (B.L. Fisher et al.) ( CAS); Reserve Forestière Majimbini, -12.76894, 45.19021, 350 m, rainforest (B.L. Fisher et al.) ( CAS).

Diagnosis.

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.

Description.

Minor worker. In full-face view, head sides diverging towards broadly convex posterior margin; eye convex and large (EL/CS: 0.26 ± 0.02; 0.21-0.30), not breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/4 of head (PoOc/CL: 0.25 ± 0.01; 0.22-0.28); frontal carinae widely diverging posteriorly (FR/CS: 0.34 ± 0.01; 0.31-0.37), distance between them larger than their smallest distance to eye; clypeus with anterolateral angle and straight anteromedian margin; mandible with two apical teeth distantly spaced; antennal scape relatively long (SL/CS: 1.23 ± 0.07; 1.01-1.36). Promesonotum slightly convex; posterior portion of mesonotum flat immediately anterior to feebly visible metanotal groove; propodeal dorsum immediately posterior to metanotal groove convex, medially strongly concave, then posteriorly flat, joining declivity at a remarkably visible angle; declivity height 3/4 length of dorsum. Petiolar node short and high, with dorsal margin inclined posteriorly and rounding to anterior margin; anterior face shorter than posterior face; femur of hind leg rounded axially, not twisted basally.

First and second gastral tergites with a pair of white spots; lateral margin of head lacking erect hairs; two erect hairs present close to posterior margin of head; antennal scape covered only with appressed hairs; pronotum with two and mesonotum with one pairs of erect hairs, respectively; posterodorsal corner of propodeum with a pair of erect hairs. Body color shining reddish black, dark brown to orange-yellow; apical portion of appendages pale in color.

Major worker. Differing from minor worker in the following characters: enlarged subquadrate head (CS: 2.44 ± 0.14; 2.19-2.66; CWb/CL: 1.04 ± 0.03; 1.00-1.08), with broadly concave posterior margin; anteromedian clypeal margin with median excision; antennal scape barely surpassing posterior cephalic margin; strongly built mandible; robust mesosoma, with distinct metanotum; propodeal dorsum slightly to strongly concave towards declivity surface; petiolar node scalelike.

Distribution and biology.

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

Discussion.

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

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 var. 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 var. invalidus are synonymyzed under Camponotus quadrimaculatus in present study.

Multivariate morphometric analysis through the NC-clustering method was applied to identify the presence of variants detected by the qualitative morphology-based study. However, NC-clustering did not reveal the existence of discrete variants. The samples from each of the variants are spread across the cluster of C. quadrimaculatus . Also, its classification success is only 97.2% because one of these samples was identified by LDA as C. gibber . This might be due to the fact that one of the variants belonging to this species is very similar to C. quadrimaculatus based on the qualitative morphology-based study.