Camponotus hovahovoides Forel, 2022

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 hovahovoides Forel stat. rev.

Figs 20B , 21B , 22A , 59

Camponotus hovahovoides Forel, 1892: 232. Syntype minor and major workers, queen and male, Madagascar, Andrangoloaka (Sikora) ( MHNG) [examined]; 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101335. Paralectotypes with same data as lectotype but: 1 major worker CASENT0101851 ( MHNG), 1 queen CASENT0101629 ( MHNG), 1 male CASENT0101759 ( MHNG) [examined]. [As subspecies of Camponotus hova Dalla Torre, 1893: 235]. Combination in Camponotus (Tanaemyrmex) : Emery, 1925: 85. Stat. rev.

Camponotus radamae var. hovoides Dalla Torre, 1893: 249. Syntype minor and major workers, queen and male, Madagascar; 1 syntype minor worker designated as lectotype, by present designation, Antananarivo ( Camboué) AntWeb CASENT0101421 ( MNHN) [examined]. Paralectotypes: 2 minor workers CASENT0101429 ( MNHN), CASENT0101337 ( MHNG); 1 queen CASENT0101412 ( MNHN); Madagascar (Hildebrandt) 1 major worker CASENT0101954 ( MHNG) and 1 male CASENT0101782 ( MHNG) [examined]. [First available use of Camponotus maculatus radamae Camponotus hovoides Forel, 1891: 33; unavailable name]. Combination in Camponotus (Tanaemyrmex) : Emery, 1925: 85. Syn. nov.

Additional material examined.

Madagascar: Antananarivo: 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (Fisher, Griswold 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); Forêt de galerie, Andranorovitra, 24.0 km NNE Ankazobe, -18.11243, 47.19757, 1491 m, disturbed gallery montane forest (B.L. Fisher et al.) ( CAS); Forêt de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, -18.12167, 47.20627, 1520 m, disturbed gallery montane forest (B.L. Fisher et al.) ( CAS); Mandraka, -18.91813, 47.91717, 1312 m, montane rainforest (B.L. Fisher et al.) ( CAS); Région Analamanga, SF Mandraka, -18.9183, 47.91687, 1285 m, montane rainforest (B.L. Fisher, F.A. Esteves et al.) ( CAS); Réserve Naturelle Sohisika, Sohisika 24.6 km NNE Ankazobe, -18.10322, 47.18692, 1464 m, gallery montane forest (B.L. Fisher et al.) ( CAS); RS Ambohitantely, -18.22444, 47.2774, 1490 m, montane forest (B.L. Fisher et al.) ( CAS); RS Ambohitantely, -18.18762, 47.28576, 1580 m, montane forest (B.L. Fisher et al.) ( 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); RS Ambohitantely, Forêt d Ambohitantely, Jardin Botanique, 24.1 km 59° NE d Ankazobe, -18.17139, 47.28182, 1620 m, montane rainforest (Fisher, Griswold et al.) ( CAS); SF Angavokely, -18.92207, 47.74157, 1460 m, rainforest (B.L. Fisher et al.) ( CAS). Antsiranana: PN Montagne d’Ambre, -12.51389, 49.17784, 984 m, montane rainforest (B.L. Fisher et al.) ( CAS); 11.0 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.45, 1585 m, montane rainforest (B.L. Fisher) ( CAS); 11.0 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.45, 1565 m, montane rainforest (B.L. Fisher) ( CAS); 11.0 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.45, 1550 m, montane rainforest (B.L. Fisher) ( CAS); 11.0 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.45, 1565 m, montane rainforest (B.L. Fisher) ( CAS); 9.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.46667, 1280 m, montane rainforest (B.L. Fisher) ( CAS); 9.2 km WSW Befingotra, Réserve Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) ( CAS); Forêt d’Ampondrabe, 26.3 km 10° NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest (B.L. Fisher) ( CAS); Forêt de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 800 m, rainforest (B.L. Fisher) ( CAS); Forêt de Binara, 9.4 km 235° SW Daraina, -13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher) ( CAS); PN Marojejy, 25.4 km 30° NNE Andapa, 10.9 km 311° NW Manantenina, -14.445, 49.735, 2000 m, montane shrubland (B.L. Fisher) ( CAS); PN Marojejy, 25.7 km 32° NNE Andapa, 10.3 km 314° NW Manantenina, -14.445, 49.74167, 1575 m, montane rainforest (B.L. Fisher) ( CAS); PN Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, -14.44333, 49.74333, 1325 m, montane rainforest (B.L. Fisher) ( CAS); RS Manongarivo 17.3 km 218° SW Antanambao, -14.02167, 48.41833, 1580-1600 m, montane rainforest (B.L. Fisher) ( CAS); RS Manongarivo, 20.4 km 219° SW Antanambao, -14.04667, 48.40167, 1860 m, montane rainforest (B.L. Fisher) ( CAS); PN Marojejy, 10.5 km NW Manantenina, -14.43333, 49.75, 1625 m, montane rainforest (E.L. Quinter) ( CAS); PN Marojejy, 11 km NW Manantenina, -14.45, 49.73333, 1875 m, montane rainforest (E.L. Quinter) ( CAS). Fianarantsoa: 28 km SSW Ambositra, Ankazomivady, -20.775, 47.16833, 1670 m, disturbed montane rainforest (B.L. Fisher) ( CAS); 29 km SSW Ambositra, Ankazomivady, -20.77667, 47.165, 1700 m, disturbed montane rainforest (B.L. Fisher) ( CAS); 28,5 km SW Ambositra, -20.78414, 47.16699, 1780 m, disturbed montane rainforest along road side (B.L. Fisher et al.) ( CAS); 45 km S Ambalavao, -22.21667, 47.01667, 785 m, rainforest (B.L. Fisher) ( CAS); 38 km S Ambalavao, PN Andringitra, -22.2, 46.96667, 1680 m, montane rainforest (B.L. Fisher) ( CAS); 40 km S Ambalavao, PN Andringitra, -22.21667, 46.96667, 1200-1275 m, rainforest (B.L. Fisher) ( CAS); 8.0 km NE Ivohibe, -22.42167, 46.89833, 1200 m, montane rainforest (B.L. Fisher, Sylvain) ( CAS); 9.0 km NE Ivohibe, -22.42667, 46.93833, 900 m, rainforest (B.L. Fisher, Sylvain) ( CAS); RS Ivohibe, 6.5 km ESE Ivohibe, -22.49667, 46.955, 1575 m, montane rainforest (B.L. Fisher, Sylvain) ( 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); Antohatsahomby II Non Protected Area, 23.38 km NW Itremo, -20.55444, 46.58438, 1640 m, Uapaca woodland (A. Ravelomanana) ( CAS); Belle Vue trail, PN Ranomafana, -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin’Hala) ( 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); Forêt d’Atsirakambiaty, 7.6 km 285 °WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) ( CAS); JIRAMA water works near river, PN Ranomafana, -21.2485, 47.45217, 690 m, open area near stream (R. Harin’Hala) ( CAS); Mampiarika IV Non Protected Area, 27.98 km SW Ambositra, -20.73528, 47.08382, 1486 m, Uapaca woodland (A. Ravelomanana) ( 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); PN Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, -23.83517, 46.96367, 940 m, rainforest (B.L. Fisher et al.) ( CAS); PN Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, -23.84083, 46.9575, 1250 m, montane rainforest (B.L. Fisher et al.) ( CAS); PN Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, -21.23667, 47.39667, 1150 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, -21.23667, 47.39667, 1150 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, -21.23667, 47.39667, 1150 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Isalo, 9.1 km 354° N Ranohira, -22.48167, 45.46167, 725 m, gallery forest (Fisher, Griswold et al.) ( CAS); Parc naturel communautaire, radio tower, PN Ranomafana, -21.25833, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M. Irwin, R. Harin’Hala) ( CAS); RS Manombo 24.5 km 228° Farafangana, -23.01583, 47.719, 30 m, rainforest (B.L. Fisher et al.) ( CAS); Vohiparara broken bridge, -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin’Hala) ( CAS); Col des Tapias, -20.26667, 47.11667, 1500 m, Uapaca woodland ( CAS); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) ( CAS). Mahajanga: Region Sofia, Bemanevika, -14.32826, 48.58406, 1657 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). Toamasina: Ambatovy, 12.4 km NE Moramanga, -18.83937, 48.30842, 1080 m, montane rainforest (B.L. Fisher et al.) ( CAS); Analamay, -18.80623, 48.33707, 1068 m, montane rainforest (Malagasy ant team) ( CAS); Ankerana, -18.40636, 48.80254, 1108 m, montane forest (B.L. Fisher et al.) ( CAS); Ankerana, -18.4104, 48.8189, 855 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80424, 48.40081, 968 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80398, 48.40358, 1064 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80398, 48.40358, 1064 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80388, 48.40506, 1013 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambohibolakely, -18.77908, 48.36628, 1014 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Ambohibolakely, -18.76087, 48.37128, 1044 m, rainforest (B.L. Fisher et al.) ( CAS); Corridor Forestier Analamay-Mantadia, Tsaravoniana, -18.75641, 48.42195, 1036 m, rainforest (B.L. Fisher et al.) ( CAS); District of Moramanga, 29 km E of Moramanga; PN Andasibe, -18.93767, 48.41167, 822 m, rainforest (Mike, Rin’ha) ( CAS); Forêt Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (Malagasy ant team) ( CAS); Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) ( CAS); Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, -15.17833, 49.635, 1100 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Andasibe-Mantadia, Forêt de Mantadia, 25.7 km 248° Moramanga, -18.81402, 48.43028, 1040 m, rainforest (F.N. Raharimalala, B. Blaimer) ( CAS); PN Zahamena, -17.73359, 48.72625, 950 m, rainforest (B.L. Fisher et al.) ( CAS); PN Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, -17.74298, 48.72936, 860 m, rainforest (B.L. Fisher et al.) ( CAS); Parcelle E3 Tampolo, -17.28104, 49.43012, 10 m, littoral forest (Malagasy ant team) ( CAS); SF Analamazaotra, Analamazaotra 1.3 km S Andasibe, -18.38466, 48.41271, 980 m, montane rainforest (B.L. Fisher et al.) ( CAS); Torotorofotsy, -18.87082, 48.34737, 1070 m, montane rainforest, marsh edge (Malagasy ant team) ( CAS). Toliara: 6 km SSW Eminiminy, Res. Andohahela, -24.73333, 46.8, 330 m, rainforest (P.S. Ward) ( CAS); Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, -24.13632, 47.05485, 1315 m, rainforest (B.L. Fisher, F.A. Esteves et al.) ( CAS); Anosy Region, Anosyenne Mts, 31.2 km NW Manantenina, -24.13894, 47.06804, 1125 m, rainforest (B.L. Fisher, F.A. Esteves et al.) ( CAS); FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) ( CAS); FC Analavelona, 33.2 km 344° NNW Mahaboboka, -22.64333, 44.17167, 1300 m, montane rainforest (Fisher, Griswold et al.) ( CAS); PN Andohahela, Manangotry, 33.8 km NW Tolagnaro, -24.75117, 46.85783, 575 m, rainforest (B.L. Fisher et al.) ( CAS); PN Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, -24.76389, 46.75167, 900 m, montane rainforest (Fisher-Griswold Arthropod Team) ( CAS); RS Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, -18.26667, 45.40667, 1050 m, montane rainforest (Fisher, Griswold et al.) ( CAS); RS Kalambatritra, Ambinanitelo, -23.4502, 46.45658, 1325 m, montane rainforest (B.L. Fisher et al.) ( CAS); RS Kalambatritra, Ampanihy, -23.4635, 46.4631, 1270 m, montane rainforest (B.L. Fisher et al.) ( CAS); RS Kalambatritra, Ampanihy, -23.463, 46.47057, 1269 m, montane rainforest (B.L. Fisher et al.) ( CAS); RS Kalambatritra, Befarara, -23.4178, 46.4478, 1390 m, montane rainforest (B.L. Fisher et al.) ( CAS); RS Kalambatritra, Betanana, -23.4144, 46.459, 1360 m, montane rainforest (B.L. Fisher et al.) ( CAS).

Diagnosis.

Lateral cephalic margins approximately parallel in full-face view; two apical teeth of mandible normally spaced; antennal scape covered with suberect hairs; in lateral view, dorsum of mesosoma from mid-mesonotum to posterodorsal corner of propodeum approximately straight, propodeal dorsum ca. 3 × as long as the height of declivity surface; petiolar node flattened anteroposteriorly.

Description.

Minor worker. Head sides anterior to level of eye parallel; lateral margins posterior to level of eye converging progressively to posterior margin; length of posterior portion of head behind eye level 1/3 length of head (PoOc/CL: 0.28 ± 0.01; 0.25-0.31). Eyes protruding and large (EL/CS: 0.30 ± 0.01; 0.27-0.34), breaking lateral cephalic margin. Frontal carinae strongly diverging posteriorly (FR/CS: 0.30 ± 0.01; 0.28-0.33); clypeus projecting into anterolateral angle and medially with blunt angle or convex. Two apical teeth of mandible distantly spaced. Antennal scape relatively long (SL/CS: 1.54 ± 0.08; 1.43-1.75) with erect to suberect hairs inclined to 45°. Promesonotum weakly convex and mesopropodeum almost flat, metanotal groove weakly visible, propodeal dorsum anteriorly convex and posteriorly flat, joining the declivity in a blunt angle, propodeal declivity height 1/2 dorsum length. Petiolar node flattened anteroposteriorly or short and high, its dorsal margin rounding to anterior margin. Tibia of hind leg rounded and not twisted dorsoventrally.

First and second gastral tergites without a pair of white spots. Lateral margin of head covered with erect hairs; posterior margin of head with two or three pairs of erect hairs; posterodorsal angle of propodeum with one or two pairs of erect hairs.

Major worker. Characteristics the same as minor worker, except the enlarged head (CS: 2.66 ± 0.20; 2.40-3.07; CWb/CL: 0.77 ± 0.02; 0.72-0.80); the more strongly built mandible; apical 1/5 of antennal scape surpassing posterior cephalic margin; metanotum visible; propodeal dorsum convex and its junction to declivity broadly angulate; petiolar node much higher than long.

Distribution and biology.

A widely distributed species, C. hovahovoides occurs mainly in the eastern lowland to montane rainforests and montane shrublands of Madagascar (Fig. 59G ). Colonies can be found mostly in rotten logs and sticks or in rotting tree stumps; they are seldom established in dead branches or twigs above the ground. Workers forage on the ground or through leaf litter, rarely on lower parts of vegetation.

Discussion.

See also discussion under C. mixtellus . Camponotus hovahovoides is one of the more common species of Myrmosaga occupying the montane rainforests of Madagascar and shows notable morphological variation. Two variants are recognized based on the shapes of the mesosoma and body colors, but these merge progressively into one typical form across the geographic distribution of the species.

Variant 1. In this variant the posterior 1/2 of mesonotum to posterodorsal corner of propodeum is not straight in lateral view and the body is pale yellow to reddish orange in color.

Variant 2. This variant is known by a dorsum of mesosoma from mid-mesonotum to posterodorsal corner of propodeum that is approximately straight, and its body is dark brown to black in color or the mesosoma is lighter in color relative to the darker head and gaster. Workers of this variant have been collected from mountaintops.

Camponotus hovahovoides has a wide distribution across the humid forest in Madagascar, this information combined with the results of the comparison of the syntype specimens of C. radamae var. hovoides with those of C. hovahovoides indicate that there is no strong morphological difference between both taxa. The few specimens of C. radamae var. hovoides represent a rare variation of C. hovahovoides across its geographic distribution.

The NC-Clustering technique was used to reveal the two variants and demonstrates that the quantitative morphological characters did not identify the difference between both variants. The members of each variant are scattered along the cluster of C. hovahovoides . More robust molecular phylogenetic information is needed to determine whether the variants constitute separate species and to study the ecological and evolutionary forces underlying these morphological variations.

Figures

Figure 20. Head in full-face view A C. mahafaly (CASENT 0115287) B C. hovahovoides (CASENT 0487242).

Figure 21. Mesosoma and petiolar node in lateral view A C. cemeryi (CASENT 0217306) B C. hovahovoides (CASENT 0496908).

Figure 22. Mesosoma in lateral view A C. hovahovoides (CASENT 0496908) B C. mixtellus (CASENT 0132606).

Figure 59. Camponotus hovahovoides A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT 0496908 and CASENT 0487242 G distribution map.