Camponotus immaculatus Forel, 1892

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 immaculatus Forel

Figs 2C , 36A , 60

Camponotus quadrimaculatus var. immaculatus Forel, 1892: 233. Syntype workers, Madagascar Andrangoloaka (Sikora); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102430 (MHNG). Paralectotypes: 2 major workers of same data as lectotype but with specimen codes: CASENT0102429, CASENT0102431 (MHNG) (Andrangoloaka). Raised to species: Olson and Ward 1996: 164.

Camponotus quadrimaculatus opacata Emery, 1925: 123. Syntype workers and alate queen, Madagascar, Baie d’Antongil (Mocquerys); 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0102107) (MSNG). Paralectotypes: 2 major worker and one alate queen of same data as lectotype but respectively with the following specimen codes: CASENT0102106 (MSNG), CASENT0102433 (MHNG) and CASENT0102108 (MSNG). [Replacement name for Camponotus quadrimaculatus opaca Emery, 1899: 290. Junior secondary homonym of Formica opaca (now Camponotus opaca ) Nylander, 1856: 55]. Syn. nov.

Additional material examined.

Madagascar: Antananarivo: [Andrangoloaka]; Mantasoa; Manjakandriana, -19.033333, 47.9166666, 1409 m (Sikora) (MHNG); Kaloy, -18.58998, 47.65102, 1423 m, disturbed montane rainforest (B.L. Fisher et al.) (CAS); Analamanga Region, District of Ankazobe, Ambohitantely, 46 km NE of Ankazobe, -18.198, 47.2815, 701 m, Forêt sclerophylle ( Rin’Ha, Mike) (CAS); Ankalalahana, -19.00711, 47.1337, 1350 m, Uapaca woodland (B.L. Fisher et al.) (CAS); Antaponimanadala I Non Protected Area, 6.59 km E Manalalondo, -19.25528, 47.1771, 1984 m, Savannah grassland (A. Ravelomanana) (CAS); Antaponimanadala II Non Protected Area, 6.63 km E Manalalondo, -19.255, 47.17684, 1955 m, Savannah grassland (A. Ravelomanana) (CAS); Navoatra I Non Protected Area, 7.64 km NW Arivonimamo, -18.97806, 47.11929, 1373 m, Uapaca woodland (A. Ravelomanana) (CAS); SF Manjakatompo, -19.35, 47.31667, 1600 m, montane rainforest (P.S. Ward) (CAS); SF Manjakatompo, -18.91667, 47.53333, 1300-1350 m, park/garden (P.S. Ward) (PSWC); 1 km NE Ambalavao, nr. Antananarivo, -19.08333, 47.53333, 1350 m, roadside (P.S. Ward) (CAS); 42 km S Antsirabe, -20.25, 47.1, 1500 m, Uapaca woodland (P.S. Ward) (CAS); Angavokely, -18.93333, 47.75 (B. Pettersson) (CAS). Antsiranana: Forêt de Binara, 9.4 km 235° SW Daraina, -13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher) (CAS); RS Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest edge (Fisher, Griswold et al.) (CAS); RS Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest, (J.-J. Rafanomezantsoa et al.) (CAS); RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, open secondary vegetation (Fisher, Griswold et al.) (CAS); PN Marojejy, 11 km NW Manantenina, -14.45, 49.73333, 1875 m, montane rainforest (E.L. Quinter) (CAS); Marojejy, tributary Manantenina River, -14.43333, 49.75, 750 m, (Quinter & Nguyen) (CAS); Galoko chain, Mont Galoko, -13.5888, 48.72864, 980 m, montane forest (B.L. Fisher et al.) (CAS). Fianarantsoa: 2 km NE Anjoma-Ramartina, -19.63333, 45.96667, 750 m, grassland (P.S. Ward) (MSNG); 28 km SSW Ambositra, Ankazomivady, -20.775, 47.16833, 1670 m, grassland (B.L. Fisher) (CAS); Ampangabe II Non Protected Area, 21.29 km W Itremo, -20.61139, 46.60809, 1402 m, savannah woodland (A. Ravelomanana) (CAS); Antapia I Non Protected Area, 26.43 km SW Ambositra, -20.71972, 47.08685, 1495 m, Uapaca woodland (A. Ravelomanana) (CAS); Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CAS); Manandriana IV Non Protected Area, 26.96 km SW Ambositra, -20.73111, 47.09512, 1594 m, savannah grassland (A. Ravelomanana) (CAS); Manandriana IV Non Protected Area, 26.96 km SW Ambositra, -20.73111, 47.09512, 1594 m, Savannah grassland (A. Ravelomanana) (CAS); Miandritsara Forest, 40 km S of Ambositra, -20.79267, 47.17567, 822 m, Low altitude 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, Plateau d’Andohariana, 39.8 km 204° Ambalavao, -22.18767, 46.90083, 2150 m, rubicole thicket at base of cliff (B.L. Fisher et al.) (CAS); Fianarantsoa, Soanierenana III Non Protected Area, 25.25 km SW Ambositra, -20.72194, 47.11019, 1707 m, savannah grassland (A. Ravelomanana) (CAS); 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); 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); Col des Tapias, -20.26667, 47.11667, 1500 m, Uapaca woodland (P.S. Ward) (PSWC), Vatovavy Fitovinany Region, District of Ifanadiana, 12 km W of Ranomafana, -21.25083, 47.40717, 1127 m, forest edge, open area ( Rin’Ha, Mike) (PSWC), Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, grassland (Fisher, Griswold et al.) (CAS); Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, grassland (Fisher, Griswold et al.) (CAS); Manakara, -22.14817, 48.02267, 10 m, urban gardens, coastal Casuarina equisetifolia (B.L. Fisher et al.) (CAS); Miandritsara Forest, 40 km S of Ambositra, -20.79267, 47.17567, 822 m, Low altitude rainforest ( Rin’Ha, Mike) (CAS); PN Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland (Fisher, Griswold et al.) (CAS); Tsaranoro, 32.8 km 230° Ambalavao, -22.08317, 46.774, 975 m, savannah woodland (B.L. Fisher et al.) (CAS). Mahajanga: PN Ankarafantsika, Ampijoroa SF, 160 km N Maevatanana, deciduous forest, -16.31944, 46.81333, 43 m, deciduous forest, Mike (Irwin, Rin’ha), Harin’Hala (CAS); PN Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on Tsingy (Fisher-Griswold Arthropod Team) (CAS); Réserve forestière Beanka, 50.7 km E Maintirano, -17.8873, 44.47113, 160 m, savannah woodland (B.L. Fisher et al.) (CAS); 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 (Mike, Rinha) (CAS). Toamasina: [Antongil]; Maroantsetra, -15.43333, 49.75, 12 m, (Mocquerys) (CAS); 6 km ESE Andasibe (=Perinet), -18.95, 48.46667, 900 m, roadside (P.S. Ward) (CAS); Brickaville, -18.82183, 49.07017, 24 m, urban/garden (B.L. Fisher et al.) (CAS); Forêt Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (B.L. Fisher) (CAS); Manakambahiny Atsinanana, -17.75, 48.71667, Primary forest (A. Pauly) (CAS); Forêt Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane shrubland, on rock (B.L. Fisher) (CAS). Toliara: Anosy Region, PN Andohahela, Forêt de Manatalinjo, -24.82466, 46.60111, 100 m, spiny forest/thicket (B.L. Fisher, F.A. Esteves et al.) (CAS); Betroka, -23.26558, 46.09716, 800 m, town/park/garden (B.L. Fisher et al.) (CAS). Ivahona, -23.45591, 46.17376, 820 m, village/park/garden (B.L. Fisher et al.) (CAS); 2 km SW Mahamavo, PN Andohahela, -24.78333, 46.7, 320 m, tropical dry forest (P.S. Ward) (MSNG); 34.6 km NW Tsihombe, -25.12983, 45.21033, 150 m, spiny forest/thicket, road side (B.L. Fisher et al.) (MHNG); 45 km NE Morondava, -20.05, 44.61667, 30 m, tropical dry forest (P.S. Ward) (CAS); 48 km ENE Morondava, -20.06667, 44.65, 30 m, tropical dry forest (D.M. Olson) (CAS); 4 km N Isaka-Ivondro, -24.76667, 46.86667, 180 m, roadside (P.S. Ward) (CAS); Anosy Region, District of Amboasary, 58 km SW of Fort Dauphin, 08 km NW of Amboasary, Berenty Special Reserve, -25.00667, 46.30333, 85 m, Galery forest ( Rin’Ha, Mike) (CAS); Anosy Region, District of Amboasary, PN Andohahela, Parcelle III, Ihazofotsy, 32 km NE Amboasary, -24.83083, 46.53617, 58 m, dry forest, spiny forest (Michael Irwin, Frank Parker, Rin’ha) (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); Anosy Region, District of Fort-Dauphin, PN Andohahela, Parcelle II, Tsimela, 42 km W of Fort-Dauphin, -24.93683, 46.62667, 176 m, transition forest (Michael Irwin, Frank Parker, Rin’ha) (CAS); Atsimo Andrefana Region, District of Betioky, 30 km E Betioky, RS Beza Mahafaly (Around Research Station), -23.6865, 44.591, 165 m, Galery dry deciduous forest ( Rin’Ha, Mike) (CAS); FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CAS); Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.045, 44.66222, 100 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); Mahafaly Plateau, 6.2 km 74° ENE Itampolo, -24.65361, 43.99667, 80 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); Makay Mts., -21.31364, 45.14782, 525 m, Gallery forest on sandy soil (B.L. Fisher et al.) (CAS); Manatantely, 8.9 km NW Tolagnaro, -24.9815, 46.92567, 100 m, rainforest (B.L. Fisher et al.) (CAS); Manderano, -23.5275, 44.08833, 70 m, gallery forest (Frontier Project) (CAS); PN Andohahela, Forêt d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, -24.93, 46.6455, 300 m, tropical dry forest (Fisher-Griswold Arthropod Team) (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 Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, -23.99222, 43.88067, 90 m, spiny forest/thicket (Fisher-Griswold Arthropod Team) (CAS); PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CAS); Portuguese Island, nr. Fort Dauphin; Ifarantsa; Taolagnaro, -24.9167, 46.8667, 30 m, lawn, D. Whitacre (CAS); RS Beza Mahafaly, Parcel 1, -23.65, 44.63333, 130 m, tropical dry forest (P.S. Ward) (CAS); Réserve Berenty, -25.01667, 46.3, 25 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); Sakaraha, -22.91233, 44.53283, 470 m, urban/garden (B.L. Fisher et al.) (CAS); Sept Lacs, -23.52083, 44.15972, 120 m, gallery forest (Frontier Project) (CAS); 27 km N Tongobory, Rupelian zone in dry forest, -23.34617, 44.35817, 290 m, spiny scrubland damp wash (M.E. Irwin, Rin’ha) (CAS); PN Andohaela Tsimelahy, -24.93683, 46.62667, 180 m, transition forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); Berenty Special Reserve, 8 km NW Amboasary, 58 km SW of Fort Dauphin, -25.00667, 46.30333, 85 m, gallery forest (M.E. Irwin, F.D. Parker, R. Harin’Hala) (CAS); PN Andohahela, Ihazofotsy - Parcel III, transition forest, -24.83483, 46.48683, 80 m, tropical dry forest, transition between spiny and dry deciduous forests (M.E. Irwin, F.D. Parker, 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).

Diagnosis.

In full-face view, lateral margins of head anterior to eye level diverging posteriorly; anterior clypeal margin truncate; white spots absent on second and third abdominal tergite.

Description.

Minor worker. With head in full-face view, lateral margins diverging towards almost straight posterior margin; eye slightly convex and small (EL/CS: 0.25 ± 0.01; 0.23-0.28), not interrupting lateral cephalic border, level of its posterior margin located at ca. posterior 1/4 of head (PoOc/CL: 0.26 ± 0.02; 0.22-0.28); frontal carinae wide, posteriorly diverging (FR/CS: 0.32 ± 0.01; 0.30-0.33), distance between them larger than their smallest distance to eye; clypeus with bluntly angulate anterolateral corner and straight anteromedian margin; two apical teeth of mandible normally distant; antennal scape relatively short (SL/CS: 1.28 ± 0.05; 1.19-1.34). Promesonotum evenly convex; posterior portion of mesonotum flat immediately anterior to weakly visible metanotal groove; propodeal dorsum strongly concave medially; junction of dorsal margin of propodeum with declivity bluntly angulate; propodeal dorsum ca. 2 × as long as declivity. Petiole nodiform, its dorsal margin inclined posteriorly and rounding to anterior margin; anterior and posterior faces almost the same height; femur of hind leg rounded axially, not twisted basally.

First and second gastral tergites without a pair of white spots; erect hairs on lateral margin of head absent; two erect hairs present near posterior cephalic margin; antennal scape with appressed hairs only; pronotum and mesonotum with a pair of erect hairs each, posterodorsal angle of propodeum with a pair of erect hairs. Integument shining; body color dark brown; distal portion of leg reddish brown.

Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 2.32 ± 0.12; 2.13-2.46; CWb/CL: 1.00 ± 0.03; 0.96-1.04), with slight concavity of posterior margin; anteromedian margin of clypeus noticeably concave; apical 1/4 of antennal scape extending beyond posterior cephalic margin; robust mesosoma, promesonotum convex, propodeal dorsum straight immediately posterior to metanotum, feebly concave towards declivity, length the same as height of declivity; petiolar node tapering dorsally.

Distribution and biology.

Occupying the high plateau of Madagascar, C. immaculatus occurs in shrubland and grassland areas, montane rainforest, savannah grassland, and Uapaca woodland habitats (Fig. 60D ). It is also known from rainforest and forest transitioning to montane forest in the southeast, and from human-modified habitats in the west, south, and high plateau of the island. Nest sites are typically established in the ground, under stones, and under rotten logs, while foraging is carried out in the leaf litter, on the ground, and on lower vegetation.

Discussion.

Camponotus immaculatus can be easily separated from C. kelimaso and C. lubbocki by the presence of the broad concavity on the propodeal dorsum. It also can be distinguished from other species because the anteromedian margin of its clypeus is truncate and no white spots are visible on the abdominal tergites.

When describing C. quadrimaculatus opacata , Emery (1925) did not make any comparative study with C. immaculatus . He stated no morphological characters that separate the former species from the later, and our examination of the syntype specimens of both taxa indicates that morphological variation observed in the former is found across the large geographical distribution of the latter. Hence, there is no doubt to make C. quadrimaculatus opacata under synonymy here.

The delimitation argument for C. immaculatus is strengthened by the congruence between the results of traditional qualitative morphology and the NC-clustering method. However, LDA produces a high 8.4% error rate in the classification of C. immaculatus , due to a misidentification of a single minor worker as C. quadrimaculatus . The high error rate is due in large part to the small range of minor worker forms of C. immaculatus available for this study.

Figures

Figure 2. Head in full-face view A C. joany (CASENT 0408908) B C. radamae (CASENT 0066777) C C. immaculatus (CASENT 0179441).

Figure 36. Head, mesosoma and petiolar node in lateral view A C. immaculatus (CASENT 0179441) B C. lubbocki (CASENT 0486998).

Figure 60. Camponotus immaculatus A lateral view B head in full-face view C dorsal view of minor worker CASENT 0082438 D distribution map.