Camponotus darwinii Forel, 1886

Rasoamanana, Nicole & Fisher, Brian L., 2022, A taxonomic revision of the Malagasy endemic subgenus Mayria of the genus Camponotus (Hymenoptera, Formicidae) based on qualitative morphology and quantitative morphometric analyses, ZooKeys 1081, pp. 137-231 : 137

Camponotus darwinii Forel

Figures 5A , 16B , 17B , 52

Camponotus darwinii Forel, 1886: 179. Lectotype minor worker, present designation, Centre de Madagascar [Ambatomanjaka, Miarinarivo, -18.766947, 46.869107, 1343 m] (Hildebrandt), AntWeb CASENT0101391 (MNHG). Paralectotypes: one alate queen and five major workers, same data as lectotype, but specimen coded respectively as CASENT0101385 (queen) andCASENT0101386 (MNHG), CASENT0104426, CASENT0104427, FOCOL2468 and FOCOL2469 (ZMHB) (examined).

Camponotus (Myrmobrachys) darwinii Forel, 1914: 270; in Camponotus (Myrmepomis) : Emery, 1920: 258; in Camponotus (Myrmopiromis) : Wheeler, 1922: 1051; Emery, 1925: 128; Bolton, 1995: 95, 131.

Camponotus darwinii var. rubropilosus Forel, 1891: 44. Lectotype major worker, present designation, Madagascar, Antananarivo ( Camboué), AntWeb CASENT0101392 (MHNG). Paralectotypes: three major workers same data as the lectotype CASENT0101393 (MHNG), CASENT0104628 (ZMHB), CASENT0101193 (NHMB).

Camponotus (Myrmopiromis) darwinii var. rubropilosus Wheeler, 1922: 1051; Emery, 1925: 128; Bolton, 1995: 121. Syn. nov.

Camponotus darwinii r. rubropilosus var. robustior Forel, 1905: 165. Lectotype minor worker, present designation, Toliara, Menabe, Beronono; Mahabo, -21.0, 45.0, 339 m (Woltzkow), AntWeb CASENT0104629 (MNHG). Paralectotypes: two major workers, same data as lectotype, but specimen coded respectively as CASENT0101195andCASENT0101380 (MHNG).

Camponotus (Myrmopiromis) darwinii r. rubropilosus var. robustior Wheeler, 1922: 1051; Emery, 1925: 128; Bolton, 1995: 121. Syn. nov.

Worker diagnosis.

Body integument matte black, finely reticulate-punctate; mesosoma and/or gastral dorsum with dense, long, and yellowish pilosity and pubescence; anterior clypeal margin with short, rounded, rectangular lobe; petiole squamiform.

Description of minor worker.

Medium size. Absolute cephalic size (CS: 1.45 ± 0.28; 1.22-1.63). In full-face view, head more quadrate (CWb/CL: 0.37 ± 0.03, 0.35-0.41); posterior margin of head almost flat, lateral margin of head slightly convex and tapering to front. Eyes circular, placed midway from the occipital corner (PoOC/CL: 0.08 ± 0.02; 0.06-0.09). Mandibles triangular with six teeth. With head in full-face view, anterior clypeal margin with a short, rounded, rectangular lobe; less convex in lateral view, not medially marginate (ClyL/GPD: 0.25 ± 0.04, 0.23-0.28). Antennal scape circular and long, surpassing the occiput by the length of two basal funiculi (SL/CS: 0.40 ± 0.04; 0.37-0.43). In lateral view, anterodorsal corner of pronotum rounded, mesosomal dorsal outline feebly and rather evenly arcuate above, dorsal face of propodeum generally longer than declivitous face (MW/ML: 0.21 ± 0.02; 0.19-0.23; MPD/ML: 0.24 ± 0.03; 0.20-0.26). In lateral view, petiolar node cuneate, anterodosum straight basally and inclined dorsally to reach the flattened posterodorsal margin. Tibia tubular, not prismatic, its extensor profile with suberect setae. Head dorsum is matte, finely reticulate-punctate with sparse punctures on malar area; mesosoma striate in lateral view; in dorsal view, petiole, mesosoma, and appendages less shiny, finely reticulate-striolate except the abdominal sternite, which is more matte. Mandible finely reticulate-punctate with sparse oblique punctures. Hairs yellowish white, orange, or tawny colored, erect hairs usually abundant, at least on gastral tergite, often on vertex, occipital portion, and thorax dorsum; petiolar node with 4 pairs of hairs on its dorsum. Shiny black, with mandible and basitarsus reddish to brownish.

Description of major worker.

Characteristics of minor workers, except: head much wider posteriorly (CS: 2.14 ± 0.26, 1.70-2.47; CWb/CL: 1.04 ± 0.06, 0.94-1.20); lateral margins convex and converging towards base of mandible. Eyes semicircular, placed dorsolaterally midway from the occipital margin (PoOC/CL: 0.22 ± 0.01, 0.19-0.24). Clypeus much more quadrate in full-face view (ClyL/GPD: 0.72 ± 0.09, 0.48-0.81), its anterior margin slightly concave medially. Antennal scape short, just surpassing the occipital border (SL/CS: 0.83 ± 0.08, 0.75-0.99). Dorsal outline of mesosoma forms a continuous line so that the basal portion of propodeum forms an obtuse angle with the declivity. Lateral portion of head finely reticulate above, punctate to finely areolate below.

Distribution and biology.

Camponotus darwinii is a famous species which is distributed widely across central of Madagascar, with a range extending from Ankazobe to Mount Papango, PN Befotaka Midongy (Fig. 58A ). It has been collected from a wide variety of different habitats, including rainforest, montane rainforest, Uapaca forest, and Uapaca woodland as well as disturbed habitats such as urban and garden areas and cultivated land (Tavy). The species has also been found nesting inside Melia azedarach and Phellolophium madagascariense . Habitat elevations ranged from 1069-2150 m.

Discussion.

Camponotus darwinii should be separable from most similar species in this group by the combination of the following characters: presence of dense hairs on mesosoma and/or gastral dorsum, mandible and malar area distinctly reddish in color, propodeal and declivitous faces unequal in length. This species is much closer to C. ursus but differs in the density of mesosomal hairs and the aspect of the petiole. In addition, C. darwinii is much more robust than C. ursus.

Additional material examined.

Province Antananarivo: Analamanga Region, District of Ankazobe, Ambohitantely , 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle (MG) (CAS); Ambatolaona, -18.928, 47.88283, 1382 m, urban/garden (BLF) (CAS); Kaloy, -18.58998, 47.65102, 1423 m, disturbed montane rainforest, (BLF) (CAS); Ilafy, -18.85415, 47.56575, 1385 m, urban/garden (BLF) (CAS); Forêt de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, -18.12167, 47.2062 7, 1520 m, disturbed gallery montane forest (BLF) (CAS); Stn. Forestiѐre Manjakatompo, -19.35, 47.31667, 2000 m, montane rainforest (Mark Pigeon) (CAS); 25 km NNE Ankazobe, -18.1, 47.18333, 1500 m, rainforest (PSW) (PSWC); Stn. Forestiѐre Manjakatompo, -19.35, 47.31667, 1600 m, montane rainforest (PSW) (PSWC); -18.91667, 47.53333, 1350 m, park/garden (PSW) (PSWC); Parc de Tsimbazaza ; Antananarivo renivohitra, -18.93, 47.52611, 1275 m (D.M. Olson) (PSWC); -18.91667, 47.53333, 1300 m, park/garden (PSW) (PSWC); 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (FGAT) (CAS); Parc Botanique et Zoologique de Tsimbazaza , -18.93233, 47.91167, 1300 m (D.H. Kavanaugh) (CAS); Angavokely, -18.93333, 47.75 (B. Pettersson) (PSWC) .

Province Fianarantsoa: 29 km SSW Ambositra, Ankazomivady , -20.77667, 47.165, 1700 m, disturbed montane rainforest (B.L. Fisher) (CAS); Forêt d’ Atsirakambiaty , 7.6 km 285° WNW Itremo , -20.59333, 46.56333, 1550 m, montane rainforest (FGAT)(CAS) ; 2 km W Andrambovato, along river Tatamaly , -21.51167, 47.41, 1075 m, cultivated land (tavy), recent 1-2 year-old tavy (BLF) (CAS); stream area , 900 m E of Isalo National Park Interpretive Center, -22.62667, 45.35817, 750 m, open area near stream (R. Harin’Hala) (CAS); Parc National d’Andringitra, Plateau d’Andohariana, 39.8 km 204° Ambalavao , -22.18767, 46.90083, 2150 m, rubicole thicket at base of cliff (BLF) (CAS); Vohiparara , -21.23333, 47.36667 (A. Pauly) (CAS); Miandritsara Forest , 40 km S of Ambositra , -20.79267, 47.17567, 822 m, low altitude rainforest (MG) (CAS); dry wash , 1 km E of Isalo National Park Interpretive Center , -22.62667, 45.35817, 885 m, dry wash (R. Harin’Hala) (CAS); Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango , -23.84083, 46.9575, 1250 m, montane rainforest (BLF) (CAS); Parc naturel communautaire , 29.3 km SW Ambositra , -20.79751, 47.1791, 1782 m, disturbed montane rainforest (BLF) (CAS); Parc naturel communautaire , 26.8 km SW Ambositra , -20.775, 47.18362, 1755 m, disturbed montane rainforest (BLF) (CAS); Parc naturel communautaire , 28.5 km SW Ambositra, -20.78414, 47.16699, 1780 m, disturbed montane rainforest along road side (BLF) (CAS); Mampiarika IV Non Protected Area , 27.98 km SW Ambositra , -20.73528, 47.08382, 1486 m, Uapaca woodland (ARA) (CAS); R.S. Ivohibe , 6.5 km ESE Ivohibe , -22.49667, 46.955, 1575 m, montane rainforest (B.L. Fisher (Sylvain )) (CAS); P. N. Andringitra, Forêt Ravaro 12.5 km SW Antanifotsy, -22.21167, 46.845, 1650 m, montane rainforest (S. Razafimandimby) (CAS); R.S. Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (B.L. Fisher (Sylvain)) (CAS); 28 km. SSW Ambositra , Ankazomivady, -20.775, 47.16833, 1670 m, montane rainforest edge (B.L. Fisher) (CAS) ; 38 km S Ambalavao, Res. Andringitra , -22.2, 46.96667, 1680 m, montane rainforest (B.L. Fisher) (CAS) ; 40 km S Ambalavao, Res. Andringitra , -22.21667, 46.96667, 1275 m, montane rainforest (B.L. Fisher) (CAS); Res. Andringitra ; 8.5 km SE Antanitotsy, -22.16667, 46.96667, 1990 m, montane rainforest (Sylvian) (CAS); 8.0 km NE Ivohibe, -22.42167, 46.89833, 1200 m, montane rainforest (B.L. Fisher (Sylvain )) (CAS); Antapia I Non Protected Area , 26.43 km SW Ambositra, -20.71972, 47.08685, 1495 m, Uapaca woodland (ARA) (CAS); Antohatsahomby I Non Protected Area , 22.77 km NW Ambatofinandrahana, -20.55056, 46.58562, 1550 m, Uapaca woodland (ARA) (CAS); Amoron’i Mania Region, District of Ambositra, Italaviana Uapaca forest , 135 Km SE of Antsirabe, -20.17333, 47.086, 1359 m, Uapaca forest, (MG) (CAS); Antapia IV Non Protected Area , 26.42 km SW Ambositra, -20.71917, 47.0868, 1494 m, Uapaca woodland (ARA) (CAS); Antohatsahomby V Non Protected Area , 22.63 km NW Itremo, -20.56722, 46.57923, 1726 m, Uapaca woodland (ARA) (CAS); Manombo Special Reserve , 32 km SE of Farafangana, -23.02183, 47.72, 36 m, lowland rainforest (MG) (CAS); Fitovavy Fitovinany Region, District of Ifanadiana , 12 km W of Ranomafana, -21.25083, 47.40717, 1127 m, forest edge, open area (MG) (CAS); Antohatsahomby IV Non Protected Area , 22.67 km NW Itremo, -20.56306, 46.58097, 1708 m, Uapaca woodland (ARA) (CAS); Ambinanindranomena Non Protected Area , 39.45 km SE Ambalavao, -21.95386, 47.29427, 1069 m, montane rainforest (ARA) (CAS); Hte Mananjary; Ambinanindrano; Ambositra , -20.733334, 47.633335, 657 m (R. Catala) (MNHN) .

Province Mahajanga: Sofia Region, District of Sofia, Anjiamangirana 45 km S Antsohihy, Analagnambe Galery forest , 5 km W Anjiamangirana, -15.157, 47.73417, 97 m, low degraded dry forest (MG) (CAS). Province Toamasina: Ambanizana, Parc National Masoala, -15.57222, 50.00694, 1020 m, montane rainforest (D. Andriamalala, D. Silva et al.) (CAS); Moramanga, -18.948902, 48.224426, 914 m (Decarpentries) (NHMB); 6.9 km NE Ambanizana, Ambohitsitondroina , -15.58506, 50.00952, 825 m, rainforest (B.L. Fisher) (CAS); Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, -15.17833, 49.635, 1100 m, montane rainforest (FGAT) (CAS). Province Toliara: [S.W. Madagascar]; Beronono; Mahabo, -21, 45, 339 m ( Voltzkow ) (NHMB); Androy Region , District of Tsihombe , 74 km S of Tsihombe, Cap Ste Marie Reserve , -25.58767, 45.163, 36 m, spiny bush (MG) (CAS); 13 km NW Enkara, Res Andohahela , -24.55, 46.8, 1300 m, montane rainforest (B.L. Fisher) (CAS); Atsimo Andrefana Region , District of Tulear II , Mikea decidious dry forest 3 km N Andranomavo village, -22.90367, 43.4755, 30 m, deciduous dry forest (MG) (CAS); Miary, -23.311972, 43.740917, 55 m (F. Geay) (MNHN); Ambovombe, -25.1775, 46.09283, 25 m (R. Decary) (MNHN); near road, Zombitse National Park , -22.8405, 44.73117, 825 m, spiny deciduous forest (R. Harin’Hala) (CAS) .

Figures

Figure 5. Morphological character of minor and major workers of C. darwinii species group A head of minor worker in full-face view of C. darwinii (CASENT 0179460) B body in lateral view of the same species C head of major worker in full-face view C. nossibeensis (CASENT 0493593) D body in lateral view of C. themistocles (CASENT 0217297).

Figure 16. Mesosomal and gastral pilosity A C. ursus (CASENT 0217284) B C. darwinii (CASENT 0071420) C C. themistocles (CASENT 0217297) D C. radovae (CASENT 0121525).

Figure 17. Dense hair on mesosoma dorsum A C. ursus (CASENT 0217284), and on gastral dorsum for B C. darwinii (CASENT 0179460).

Figure 52. Camponotus darwinii minor worker (CASENT 0179460) 1 body in lateral view 2 head in full-face view 3 body in dorsal view.

Figure 58. Distribution map of C. darwinii species group A C. darwinii B C. norvigi C C. nossibeensis D C. radovae E C. themistocles F C. ursus