Camponotus strangulatus Santschi, 1911

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

Camponotus strangulatus Santschi

Figs 29A , 74

Camponotus maculatus st. strangulatus Santschi, 1911a: 129. Lectotype major worker, by present designation, Madagascar, Vitikampy; Morondava, AntWeb CASENT0101097 (NHMB) [examined]. Raised to species by Olson and Ward 1996: 164.

Camponotus hova maculatoides Emery, 1920b: 6. Lectotype minor worker, by present designation, Madagascar, Nosibe, Antsiranana (Voeltzkow) AntWeb CASENT0101095 (NHMB) [examined]. Paralectotypes: 1 minor worker and 3 major workers of same data as lectotype, but minor and majors respectively specimen coded as: CASENT0101339, CASENT0101338, CASENT0101747 (MHNG) and CASENT0101095 (NHMB) [examined]. [First available use of Camponotus maculatus r. hova var. maculatoides Forel, 1897: 200]. Syn. nov.

Camponotus strangulatus Note: As junior synonym by Emery 1920: 6. [Emery established Camponotus strangulatus as the junior synonym of Camponotus maculatoides but the former name has priority: Bolton 1995: 125].

Additional material examined.

Comoros: Grande Comore: Domani, -11.51778, 43.28, 25 m, coastal scrub (B.L. Fisher et al.) (CAS); Pidjani, -11.75447, 43.45148, 35 m, coastal scrub (B.L. Fisher et al.) (CAS). Mohéli: Madahali 50, -12.37421, 43.86857, 50 m, coastal dry forest scrub (B.L. Fisher et al.) (CAS); Ouallah, -12.34234, 43.66851, 1 m, mangrove (B.L. Fisher et al.) (CAS). Juan De Nova Island: -17.04891, 42.72102, 10 m, littoral vegetation (B.L. Fisher) (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); RS Ambohitantely, Forêt d’Ambohitantely, 20.9 km 72° NE d’Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CAS). Antsiranana: [ Loucoubé auf Nossi-Bé]; Nosy be; Nosibe (Carl Bosse) (MHNG); District of Vohemar, Analabe Sahaka 43 km E of Daraina, -13.07933, 49.90233, 30 m, wooded grassland-bushland (Mike, Rin’ha) (CAS); Forêt d’Orangea, 3.6 km 128° SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) (CAS); Galoko chain, Mont Kalabenono, -13.64179, 48.67282, 643 m, rainforest (B.L. Fisher et al.) (CAS); Galoko chain, Mont Kalabenono, -13.63999, 48.67374, 498 m, rainforest (B.L. Fisher et al.) (CAS); Nosy Be, 4 km ESE Andoany (=Hellville), -13.41667, 48.3, <5 m, littoral vegetation (P.S. Ward) (CAS); Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CAS); PN Montagne d’Ambre, Fozalanana, -12.46637, 49.22157, 475 m, dry forest (B.L. Fisher et al.) (CAS); Réserve Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) (CAS); SAVA Region District of Vohemar, Andranotsimaty, Dry forest, 9 km NE of Daraina, -13.1695, 49.70067, 90 m, dry dense forest (Mike, Rinha) (CAS); Taizambato, 22.9 km NE Ambanja, -13.5092, 48.56722, 16 m, coconut plantation (B.L. Fisher et al.) (CAS); PN Montagne d’Ambre [1st campsite], 960 m, rainforest (R. Harin’Hala) (CAS); PN Montagne d’Ambre [Petit Lac road], -12.533333, 49.166667, 1125 m, rainforest (R. Harin’Hala) (CAS). Fianarantsoa: 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); Atsimo Atsinanana Region, District of Farafangana, Mahabo Mananivo, 50 km S of Farafangana Ampitavananima, Forest low altitude, Littoral forest on sand 2 km E of Mobot office, -23.12983, 47.717, 33 m, Littoral Low Alt Rain Forest (Mike, Frank, Rin’ha) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana, 12 km W of Ranomafana, -21.25083, 47.40717, 1127 m, forest edge, open area ( Rin’Ha, Mike) (CAS); RS Manombo, 32 km SE of Farafangana, -23.02183, 47.72, 36 m, Lowland rainforest ( Rin’Ha, Mike) (CAS). Miandritsara Forest, 40 km S of Ambositra, -20.79267, 47.17567, 822 m, Low altitude rainforest ( Rin’Ha, Mike) (CAS); PN Andringitra, Forêt Ravaro 12.5 km SW Antanifotsy, -22.21167, 46.845, 1500-1800 m, montane rainforest, (S. Razafimandimby) (CAS); radio tower, PN Ranomafana, -21.25833, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M. Irwin, R. Harin’Hala) (CAS); Vohiparara broken bridge, -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin’Hala) (CAS). Mahajanga: Ambalahonko, 5.69 km S Antsohihy, -14.8439, 48.01421, 21 m, urban garden (B.L. Fisher et al.) (CAS); Boeny Region, District of Soalala, Namoroka village, Befatika Andranovory 7 km NW of Vilanandro village, -16.46967, 45.39133, 120 m, Dense dry forest (Irwin, Rin’ha) (CAS); Boeny Region: PN Ankarafantsika, Ambodimanga, -16.32342, 46.82443, 75 m, dry forest, Canopy Bootcamp, (A. Karunakaran) (CAS); Boeny Region; District of Soalala, Namoroka 53 km of Soalala, Ambatofolaka dry forest 3 km N of Vilanandro villlage, -16.47333, 45.39133, 105 m, dense dry forest in the mud (Mike, Rin’ha) (CAS); Forêt de Tsimembo, 8.7 km 336° NNW Soatana, -19.02139, 44.44067, 20 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Melaky Region, District of Maintirano, Asondrodava dry forest against dune 15 km N of Maintirano, -17.96533, 44.0355, 16 m, dry forest (Irwin, Rinha) (CAS); Melaky Region, District of Besalampy, Marofototra dry forest, 17 km W of Besalampy, -16.72167, 44.42367, 51 m, dry wash in the dry forest (Irwin, Rin’ha) (CAS); Melaky Region, District of Besalampy, Analangidro dry forest, 07 km NE of Besalampy, -16.6915, 44.5235, 25 m, dry forest on sand (Irwin, Rin’ha) (CAS); Melaky Region, District of Maintirano, Asondrodava dry forest, 15 km N of Maintirano, -17.96533, 44.0355, 6 m, dry forest (Irwin, Rinha) (CAS); Melaky Region, District of Mintirano, Ampasy 50 km E of Maintirano, -18.004, 44.452, 85 m, dry forest (Mike, Rin’ha) (CAS); Melaky Region; District of Besalampy marofototra palm forest, 17 km W of Besalampy, -16.72167, 44.42367, 10 m, Palm trees on sand (Irwin, Rin’ha) (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 Baie de Baly, 12.4 km 337° NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Namoroka, 9.8 km 300° WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest (Fisher, Griswold et al.) (CAS); PN Tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River, -19.13222, 44.81467, 100 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (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 forestière Beanka, 50.2 km E Maintirano, -17.88756, 44.47265, 153 m, tropical dry forest on tsingy (B.L. Fisher 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). Toliara: [Vitikampy; Morondava; Museum Paris, G. Grandidier 1899]; Morondava, -20.290419, 44.299996, 6 m (CAS); 48 km ENE Morondava, Kirindy, -20.06667, 44.65, 30 m, tropical dry forest (D.M. Olson) (CAS); Ambovombe, -25.1775, 46.09283, 25 m, urban/garden (B.L. Fisher et al.) (CAS); Ampanihy; Behompy, -23.209835, 43.9457622, 268 m, urban/garden (B.L. Fisher et al.) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.021, 46.3055, 36 m, spiny forest (Mike, Rin’ha) (CAS); Atsimo Andrefana Region, District of Tulear II, Mikea deciduous dry forest 3 km N Andranomavo village, -22.90367, 43.4755, 30 m, Deciduous dry forest ( Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Tulear II, Mikea spiny forest 8 km N Andranomavo village, -22.91333, 43.39883, 37 m, spiny forest ( Rin’Ha, Mike) (CAS); Atsimo Andrefana Region, District of Tulear II, Private dry forest, Ifaty 30 km N of Tulear, -23.17967, 43.61683, 9 m, vegetation on sandy area (Mike, Rin’ha) (CAS); Betioky, -23.72117, 44.38017, 270 m, urban/garden (B.L. Fisher et al.) (CAS); Beza-Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CAS); Col du Manangotry, -24.75, 46.8, 600 m, rainforest edge (B.L. Fisher) (CAS); Ehazoara Canyon, 26 km E Betioky, -23.68333, 44.63333, 175 m, tropical dry forest (B.L. Fisher) (CAS); Ejeda, -24.3505, 44.516, 250 m, urban/garden (B.L. Fisher et al.) (CAS); Fiherenana, -23.23528, 43.87083, 50 m, degraded gallery forest (Frontier Project) (CAS); Fiherenana, -23.22252, 43.88088, 65 m, gallery forest, degraded (Frontier Wilderness Project) (CAS); Forêt de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Mahavelo, Isantoria River, -24.75833, 46.15717, 110 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Forêt de Mite, 20.7 km 29° WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest (Fisher-Griswold Arthropod Team) (CAS); Forêt de Tsinjoriaky, 6.2 km 84° E Tsifota, -22.80222, 43.42067, 70 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Kirindy Forest, plot I, -20.07434, 44.67247, 60 m, dry forest (E. Lokensgard) (CAS); Mahafaly Plateau, 6.2 km 74° ENE Itampolo, -24.65361, 43.99667, 80 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Mahafaly, near Eloeste, By Lac Tsimanampetsoa, -24.16667, 43.75, (V. & B. Roth) (CAS); Manderano, -23.5275, 44.08833, 70 m, gallery forest (Frontier Project) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza dry forest 07,5 km E of Beroboka, -19.9775, 44.66633, 50 m, dry forest (M. Irwin, Rin’ha) (CAS); Menabe Region, District of Morondava, Beroboka village 45 km NE of Morondava, Antsarongaza galery forest 07 km E of Beroboka, -19.9775, 44.66533, 45 m, Galery forest (M. Irwin, Rin’ha) (CAS); PN Andohahela, Forêt de Manatalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Kirindy Mite, 16.3 km 127° SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, 6.7 km 130° SE Efoetse, 23.0 km 175° S Beheloka, -24.10056, 43.76, 25 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Tsimanampetsotsa, Mitoho Cave, 6.4 km 77° ENE Efoetse, 17.4 km 170° S Beheloka, -24.04722, 43.75317, 40 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Ranobe, -23.04055, 43.6096, 30 m, spiny forest/thicket (Frontier Project) (CAS); Ranobe, -23.04067, 43.60973, 20 m, gallery forest, degraded (Frontier Wilderness Project) (CAS); Ranobe, -23.03883, 43.60982, 30 m, gallery forest (Frontier Wilderness Project) (CAS); Ranobe, -23.03952, 43.61015, 20 m, gallery forest (Frontier Wilderness Project) (CAS); Ranobe, -23.0342, 43.61185, 30 m, spiny forest/thicket (Frontier Project) (CAS); Ranobe, -23.04457, 43.61532, 20 m, spiny forest/thicket (Frontier Wilderness Project) (CAS); RS Beza-Mahafaly, Parcel 1, -23.65, 44.63333, 130 m, tropical dry forest (P.S. Ward) (CAS); Réserve Privé Berenty, Forêt de Bealoka, Mandraré River, 14.6 km 329° NNW Amboasary, -24.95694, 46.2715, 35 m, gallery forest (Fisher-Griswold Arthropod Team) (CAS); Sept Lacs, -23.5275, 44.15444, 70 m, Gallery forest (Frontier Project) (CAS); 3 km E Itampolo, malaise across path of lower bench of Andrimpano Forest, -24.65783, 43.95617, 45 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km E Itampolo, malaise across path of plateau of Andrimpano Forest, -24.65033, 43.96317, 130 m, dry forest (M.E. Irwin, Rin’ha) (CAS); 5 km N Ampotaka, malaise on trail in Vitambany gallery forest, -24.65033, 43.96317, 86 m, Gallery forest (M.E. Irwin, Rin’ha) (CAS); Ambohimahavelona village 33 km NE of Tulear, Andoharano dry forest, -23.44083, 43.89967, 46 m, dry forest (M.E. Irwin, Rin’ha) (CAS); Ambohimahavelona village 33 km NE of Tulear, private spiny bush, -23.44083, 43.89967, 43 m, spiny bush (M.E. Irwin, Rin’ha) (CAS). Andranovato, Euphorbia forest 5 km SE of Manombo, -22.81806, 43.50217, 18 m, Ephorbia forest (CAS); Atsimo Andrefana Region, District of Tulear II,Tsifota 20 km N of Manombo, -22.818, 43.37267, 15 m, spiny forest (M.E. Irwin, Rin’ha) (CAS); Itampolo, Sud A Sud Hotel, malaise in dune vegetation, -24.6905, 43.944, 12 m, littoral bush (M.E. Irwin, Rin’ha) (CAS); Mikea Forest, deciduous dry forest, -22.90367, 43.4755, 30 m, deciduous dry forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Mikea Forest, spiny forest, -22.91333, 43.48222, 37 m, spiny forest (R. Harin’Hala) (CAS); Parcel I, RS Beza Mahafaly, near research station, -23.6865, 44.591, 165 m, dry deciduous forest (R. Harin’Hala) (CAS); Parcel II, RS Beza Mahafaly, near Bellevue, -23.68983, 44.5755, 180 m, spiny forest (R. Harin’Hala) (CAS); PN Tsimanampetsotsa, Mitoho Forest, malaise across trail at escarpment base, -24.0485, 43.75233, 120 m, dense dry forest (M.E. Irwin, Rin’ha) (CAS); Tsimelahy - Parcel II, PN Andohahela, transition forest, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS). Mayotte: Baie de Tsingoni, -12.7926, 45.10764, 5 m, mangrove, coastal scrub (B.L. Fisher et al.) (CAS); Coconi, DAF Campus, -12.83333, 45.13333 (R. Jocqué & G. DeSmet) (CAS); Tanaraki, -12.75754, 45.0678, 10 m, coastal scrub (B.L. Fisher et al.) (CAS). Seychelles: Aldabra Atoll, Grande Terre Island, -9.43453, 46.45767, 7 m, Takamaka forest, S.M. Goodman (CAS); Aldabra Island, Picard, old settlement, -9.39606, 46.20465, 2 m, coastal scrub (B.L. Fisher) (NHMB).

Diagnosis.

With head in full-face view, lateral cephalic margins anterior to eye level parallel, lacking erect hairs; in oblique profile, three pairs of erect hairs arranged successively from level of anterior ocular margin towards posterior cephalic margin; clypeus with distinct anterolateral corner; in profile, junction of propodeal dorsum to declivity rounded; petiole nodelike and not anteroposteriorly compressed.

Description.

Minor worker. In full-face view, head sides anterior to eye level parallel, converging strongly towards posterior margin behind eye level; eye convex (EL/CS: 0.28 ± 0.01; 0.26-0.30), breaking lateral cephalic margin, level of its posterior margin located approximately at posterior 1/3 of head (PoOc/CL: 0.27 ± 0.01; 0.26-0.29); frontal carinae close, distance between them equal to or smaller than their smallest distance to eye (FR/CS: 0.25 ± 0.01; 0.24-0.27); clypeus with anterolateral angle and an almost straight or bluntly angulate anteromedian margin; two apical teeth of mandible normally spaced; antennal scape relatively long (SL/CS: 1.58 ± 0.05; 1.50-1.66). Mesosoma low and long (MPH/ML: 0.33 ± 0.01; 0.31-0.36); promesonotum weakly convex, mesonotum and propodeum nearly flat, separated by weakly visible metanotal groove; propodeal dorsum rounding to declivity; propodeal dorsum roughly 2 × as long as declivity. Petiolar node tapering dorsally, its dorsal margin inclined posteriorly, rounding to anterior margin; anterior face 1/2 height of posterior face; femur of hind leg rounded axially, not twisted near base.

First and second gastral tergites without a pair of white spots; lateral margin of head without erect hairs; a pair of erect hairs present near posterior cephalic margin; with head in profile, three pairs of erect hairs arranged from level of anterior margin of eye to posterior cephalic margin; antennal scape without erect hairs, covered with distantly spaced pubescence; pronotum with two pairs of erect hairs; mesonotum and propodeum without erect hairs.

Major worker. With characteristics of minor worker, except for the following divergent features: enlarged head (CS: 3.22 ± 0.36; 2.73-3.74; CWb/CL: 0.91 ± 0.06; 0.80-0.98), apical 1/4 of antennal scape surpassing posterior cephalic margin, much more robust mesosoma, promesonotum forming a convexity, separated from propodeum by a distinct metanotum; propodeal dorsum convex immediately behind metanotum, length almost equal to height of declivity, their junction at a rounded angle; petiolar node tapering dorsally.

Distribution and biology.

Endemic to the Malagasy region, C. strangulatus occurs in Grand Comore and Moheli of the Comoros islands, in Mayotte, in Aldabra of the Seychelles, and in Madagascar (Fig. 74D ). In the Comoros and Mayotte this species occupies mangrove and coastal dry forest scrub, while in the Seychelles it also inhabits forest habitats. Regarding the distribution of C. strangulatus in Madagascar, it is found in rainforest and littoral rainforest in the north, in dry forest in the west, in spiny forest and thicket in the south, in montane rainforest and Uapaca woodland on the south-central high plateau, and in human-modified habitats in its southern range. It also occurs in littoral vegetation on Juan de Nova Island. Due to its capacity to colonize different types of habitats, C. strangulatus also builds nests in a large array of microhabitats such as rotten logs and rotten sticks on the ground, dead branches and dead twigs above the ground, under stones, and in the ground. Workers forage on the forest floor, in leaf litter, and on lower vegetation.

Discussion.

Camponotus strangulatus may be difficult to distinguish from C. tapia and C. atimo but the dorsum of the head in the latter species bears four or more pairs of erect hairs arranged successively from the level of the anterior ocular margin towards the posterior cephalic margin.

As generally known, the genus Camponotus displays polymorphism in the worker castes and this feature is amplified by the phenotypic variation within the castes when the species has specifically a large geographic distribution. This is especially true for C. strangulatus which inhabits most of the terrestrial landscapes, but not the rainforests in the east of Madagascar. Camponotus hova maculatoides , created by Emery (1920b) was found in Nosibe, north of Madagascar, an area within the range of C. strangulatus . Observations of the type specimens of the species and subspecies prove the absence of strong morphological traits to differentiate them. The comparison of the type specimens of C. hova maculatoides with the samples of workers for C. strangulatus did not show striking morphological difference. Consequently, C. hova maculatoides is placed under synonymy here.

Species delimitation of C. strangulatus on the basis of traditional taxonomic study is congruent with the results of the NC-clustering method. The species was also classified correctly at 100% by confirmatory LDA.

Figures

Figure 29. Head in lateral view A C. strangulatus (CASENT 0136581) B C. tapia (CASENT 0493939).

Figure 74. Camponotus strangulatus A lateral view B head in full-face view C dorsal view of minor worker CASENT 0132660 D distribution map.