Revision of the Malagasy Camponotus subgenus Myrmosaga (Hymenoptera, Formicidae) using qualitative and quantitative morphology Author Rakotonirina, Jean Claude Madagascar Biodiversity Center, BP 6257, Parc Botanique et Zoologique de Tsimbazaza, Antananarivo, Madagascar & Departement d'Entomologie-Culture, Elevage, Sante; Faculte des Sciences, BP 906, Universite d'Antananarivo, Antananarivo, Madagascar jcrakoto25@yahoo.com Author Fisher, Brian L. https://orcid.org/0000-0002-4653-3270 Entomology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118, USA text ZooKeys 2022 2022-05-03 1098 1 180 http://dx.doi.org/10.3897/zookeys.1098.73223 journal article http://dx.doi.org/10.3897/zookeys.1098.73223 1313-2970-1098-1 B4F4033F296E43CCBE54B9413BC19268 B22184E75B1A59C9B985C0DD9F092196 Camponotus dufouri Forel Figs 8B , 12A , 52 Camponotus dufouri Forel, 1891: 16, 74 (Key). Syntype workers, Madagascar, 30 miles NW Tamatave ( O'Swald ) (MHNG) [examined]; 1 syntype major worker designated as lectotype, by present designation, Madagascar, AntWeb CASENT0101644 (MHNG) [examined]. Paralectoype: 1 major worker same data as lectotype but with specimen code: CASENT0101669 (MHNG) [examined]. [Combination in Camponotus (Dinomyrmex) : Forel, 1914: 268; in Camponotus (Tanaemyrmex) : Emery, 1925: 85]. Camponotus dufouri imerinensis Forel, 1891: 18. Syntype workers and queen, Madagascar, Environ d'Antananarivo , Imerina [labeled as "Andrangoloaka" ] (Sikora) (MHNG) [examined]; 1 syntype minor worker designated as lectotype, by present designation, AntWeb CASENT0101812 (Sikora) (MHNG) [examined]. Paralectotypes: 1 major worker and 1 alate queen of the same data as lectotype but with the following specimen codes: CASENT0101919 (Sikora) and CASENT0101679 ( Camboue ) (MHNG) [examined]. Syn. nov. Additional material examined. Madagascar : Antananarivo : [Madagascar], Ambatomanjaka; Miarinarivo, -18.766947 , 46.869107 , 1343 m (Sicora) (CAS); 3 km 41 °NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m , montane rainforest (Fisher, Griswold et al.) (CAS); Region Analamanga, SF Mandraka, -18.9183 , 47.91687 , 1285 m , montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); [Andrangoloaca]; Mantasoa; Manjakandriana, -19.033333 , 47.9166666 , 1409 m (MHNG). Antsiranana : 9.2 km WSW Befingotra, Reserve Anjanaharibe-Sud, -14.75, 49.46667, 1280 m , montane rainforest (B.L. Fisher) (CAS); 9.2 km WSW Befingotra, Reserve Anjanaharibe-Sud, -14.75, 49.46667, 1200 m , montane rainforest (B.L. Fisher) (CAS); Binara Forest, -13.26392 , 49.59919 , 1065 m , rainforest (B.L. Fisher et al.) (CAS); Binara Forest, -13.26388 , 49.60141 , 900 m , rainforest (B.L. Fisher et al.) (CAS); Foret Ambanitaza, 26.1 km 347° Antalaha, -14.67933 , 50.18367 , 240 m , rainforest (B.L. Fisher) (CAS); Foret de Binara, 9.1 km 233° SW Daraina, -13.26333 , 49.60333 , 650-800 m , rainforest (B.L. Fisher) (CAS); Foret de Binara, 9.1 km 233° SW Daraina, -13.26333 , 49.60333 , 800 m , rainforest (B.L. Fisher) (CAS); Makirovana forest, -14.16044 , 49.95216 , 550 m , rainforest (B.L. Fisher et al.) (CAS); PN Masoala, -15.32331 , 50.30751 , 60 m , rainforest (B.L. Fisher et al.) (CAS); PN Montagne d'Ambre , Roussette Camp 7 km SW Park entrance, -12.51444 , 49.18139 , 960 m , rainforest (Mike, Frank, Rin'ha ) (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); PN Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327 °NNW Manantenina, -14.435, 49.76, 775 m , rainforest (B.L. Fisher et al.) (CAS); PN Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333 °NNW Manantenina, -14.43667 , 49.775 , 450 m, rainforest (B.L. Fisher) (CAS); PN Montagne d'Ambre , 3.6 km 235° SW Joffreville, -12.53444 , 49.1795 , 925 m , montane rainforest (Fisher, Griswold et al.) (CAS); PN Montagne d'Ambre , Antomboka, -12.51269 , 49.17807 , 970 m , montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d'Ambre , Crete , -12.58132 , 49.13368 , 1110 m , montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d'Ambre , Petit lac, -12.53664 , 49.17412 , 1130 m , montane rainforest (B.L. Fisher et al.) (CAS); PN Montagne d'Ambre , Roussettes, -12.52574 , 49.17238 , 1025 m , montane rainforest (B.L. Fisher et al.) (CAS); R.N.I. Marojejy, 11 km NW Manantenina, -14.43333, 49.75, 1225 m , montane rainforest (E.L. Quinter) (CAS); RS Manongarivo, 10.8 km 229 °SW Antanambao, -13.96167 , 48.43333 , 400 m , rainforest (B.L. Fisher) (CAS); RS Manongarivo, 12.8 km 228° SW Antanambao, -13.97667 , 48.42333 , 780 m , rainforest (B.L. Fisher) (CAS); RS Manongarivo, 14.5 km 220° SW Antanambao, -13.99833 , 48.42833 , 1175 m , montane rainforest (B.L. Fisher) (CAS); PN Marojejy, 10 km NW Manantenina, -14.43333 , 49.76667 , 750 m , rainforest (E.L. Quinter) (CAS); PN Marojejy, 8 km NW Manantenina, -14.43333 , 49.78333 , 450 m , rainforest (E.L. Quinter) (CAS); Sakalava Beach [vegetated beach dunes], -12.26972 , 49.39167 , 10 m , across sandy trail in dwarf litoral forest (R. Harin'Hala ) (CAS); SAVA Region, District of Sambava, PN Marojejy, 5 km W of Manantenina village, 1st Camp site (Mantella), -14.43817 , 49.774 , 487 m, Low altitude rainforest ( Rin'Ha , Mike) (CAS); Montaigne Francais, 150 m , along forested limestone ridge (R. Harin'Hala ) (CAS); Montaigne Francais, 150 m , along forested limestone ridge (R. Harin'Hala ) (CAS); PN Montagne d'Ambre [1st campsite], 960 m , rainforest (Irwin, Schlinger, Harin'H ) (CAS); PN Montagne d'Ambre [lemur trail], 975 m , rainforest (R. Harin'Hala ) (CAS); PN Montagne d'Ambre [Petit Lac road], -12.533333 , 49.166667 , 1125 m , rainforest (Irwin, Schlinger, Harin'H ) (CAS). Fianarantsoa : 40 km S Ambalavao, PN Andringitra, -22.21667 , 46.96667 , 1275 m , montane rainforest (B.L. Fisher) (CAS); 45 km S Ambalavao, -22.21667 , 47.01667 , 720 m , rainforest edge (B.L. Fisher) (MHNG); Andrambovato along river Tatamaly, -21.51082 , 47.40992 , 1063 m , rainforest (B.L. Fisher et al.) (CAS); Belle Vue trail, PN Ranomafana, -21.2665 , 47.42017 , 1020 m , mixed tropical forest (M.E. Irwin, F.D. Parker (R. Harin'Hala ) (CAS); Vatovavy Fitovinany Region, District of Ifanadiana Belle vue area 1.2 km S of PN Ranomafana entrance, -21.2665 , 47.42017 , 1018 m , rainforest ( Rin'Ha , Mike) (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); Foret d'Ambalagoavy Nord, Ikongo, Ambatombe, -21.857068 , 47.37849 , 625 m (R. Harin'Hala & M.E. Irwin) (CAS); Foret de Vevembe, 66.6 km 293° Farafangana, -22.791 , 47.18183 , 600 m , rainforest, transition to montane forest (B.L. Fisher et al.) (CAS); JIRAMA water works near river, PN Ranomafana, -21.2485 , 47.45217 , 690 m , open area near stream (R. Harin'Hala ) (CAS); PN Ranomafana; Ranomafana; Ifanadiana, -21.2161494 , 47.4565003 , 1087 m (B. Pettersson) (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 Ranomafana, Talatakely, -21.24833 , 47.42667 (CE Griswold, DH Kavanaugh, ND Penny, MJ Raherilalao, JS Ranorianarisoa, J Schweickert) (CAS); RS Ivohibe 8.0 km E Ivohibe, -22.48333 , 46.96833 , 1200 m , montane rainforest (B.L. Fisher, Sylvain) (CAS); RS Ivohibe, 7.5 km ENE Ivohibe, -22.47, 46.96, 900 m , rainforest (B.L. Fisher, Sylvain) (CAS); radio tower, PN Ranomafana, -21.25833 , 47.40717 , 1130 m , forest edge, mixed tropical forest, open area (M. Irwin, R. Harin'Hala ) (CAS); PN Ranomafana, Talatakely area, 0.4 km WSW of Park Entrance, -21.41667 , 47.68333 , 900 m , mixed tropical forest (D.H. & K.M. Kavanaugh) (CAS); Reserve Speciale 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). Toamasina : [Bois 30 miles NW. Tamatave], Ambodiriana; Toamasina Rural, -17.88832 , 49.22849 , 373 m (Swald) (CAS); [Tamatave?], Andranobolahy, Toamasina Rural, -18, 49, 370 m (H. O'Swald ) (CAS); 5.3 km SSE Ambanizana, Andranobe, -15.66667 , 49.96667 , 600 m , rainforest (B.L. Fisher) (MHNG); 5.3 km SSE Ambanizana, Andranobe, -15.67133 , 49.97395 , 425 m , rainforest (B.L. Fisher) (MHNG); 6.2 km SSE Ambanizana, Be Dinta, -15.66667 , 49.99806 , 600 m , rainforest (V. Razafimahatratra.) (MNHN); 6.3 km S Ambanizana, Andranobe, -15.6813 , 49.958 , 25 m, rainforest (B.L. Fisher) (MNHN); 6.9 km NE Ambanizana, Ambohitsitondroina, -15.58506 , 50.00952 , 825 m , rainforest (B.L. Fisher) (MNHN); 7 km SE PN Andasibe Headquarters, -18.969856 , 48.465894 , 1050 m , tropical forest (M.E. Irwin R. Harin'Hala ) (CAS); Analanjirofo Region, District of ToamasinA Mobot Site, Analalava humid dense forest low altitude on the sand 7 km SW of Foulpointe, -17.69333 , 49.46028 , 75 m , dense humide low alt on the sandy soil (Mike, Rin'ha ) (CAS); PN Andasibe, botanic garden near entrance, West of ANGAP office, -18.925172 , 48.418651 , 1025 m , tropical forest (M.E. Irwin R. Harin'Hala ) (CAS); Ankerana, -18.40829 , 48.82107 , 750 m , rainforest (B.L. Fisher et al.) (CAS); Antsianaka; Ambatosoratra; Ambatondrazaka, -17.58333, 48.5, 752 m (Perrot Freres) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.80398 , 48.40358 , 1064 m , rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Ambatoharanana, -18.79944 , 48.40375 , 1016 m , rainforest (B.L. Fisher et al.) (CAS); Corridor Forestier Analamay-Mantadia, Tsaravoniana, -18.76124 , 48.42134 , 939 m , rainforest (B.L. Fisher et al.) (CAS); FC Andriantantely, -18.695 , 48.81333 , 530 m , rainforest (H.J. Ratsirarson) (CAS); FC Didy, -18.19833 , 48.57833 , 960 m , rainforest (H.J. Ratsirarson) (CAS); FC Sandranantitra, -18.04833 , 49.09167 , 450 m , rainforest (H.J. Ratsirarson) (CAS); Foret Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m , montane rainforest (Malagasy ant team) (CAS); Ile Sainte Marie, Foret Ampanihy, 14.4 km 52° Ambodifotatra, -16.91117 , 49.93917 , 10 m , littoral rainforest (B.L. Fisher et al.) (CAS); Mahavelona (Foulpointe), -17.66667, 49.5 in sandy forest (A. Pauly) (CAS); Manakambahiny Atsinanana, -17.75, 48.71667 Primary forest (A. Pauly) (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 , 18.0 km 21° NNE Ambinanitelo, -15.18833 , 49.615 , 470 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 Mantadia, -18.79167 , 48.42667 , 895 m , rainforest (H.J. Ratsirarson) (CAS); PN Zahamena, Besaky River, -17.75244 , 48.85321 , 760 m , rainforest (B.L. Fisher et al.) (CAS); PN Zahamena, Onibe River, -17.75908 , 48.85468 , 780 m , rainforest (B.L. Fisher et al.) (CAS); PN Mananara-Nord, 7.1 km 261° Antanambe, -16.455 , 49.7875 , 225 m , rainforest (B.L. Fisher et al.) (CAS); Parcelle K9 Tampolo, -17.175 , 49.268 , 10 m, littoral forest (Malagasy ant team) (CAS); RNI Betampona, Camp Vohitsivalana, 37.1 km 338° -17.88667 , 49.2025 , 520 m , rainforest (B.L. Fisher et al.) (CAS); RNI Betampona, 34.08 km 332° Toamasina -17.91977 , 49.20039 , 525 m , rainforest (B.L. Fisher) (CAS); RNI Betampona, 34.1 km 332° Toamasina -17.91614 , 49.20185 , 550 m , rainforest (B.L. Fisher) (CAS); SF Tampolo, 10 km NNE Fenoarivo Atn. -17.2825, 49.43, 10 m , littoral rainforest (B.L. Fisher) (CAS); Sahafina forest 11.4 km W Brickaville, -18.81445 , 48.96205 , 140 m , rainforest (B.L. Fisher et al.) (CAS); Tamatave; Andranobolahy; Toamasina Rural, -18, 49, 370 m (CAS). Toliara : 10 km NW Enakara, PN Andohahela, -24.56667 , 46.81667 , 420 m , rainforest (B.L. Fisher) (PSWC); 13 km NW Enakara, PN Andohahela, -24.55, 46.8, 1250 m , montane rainforest (B.L. Fisher) (CAS); Anosy Region, Anosyenne Mts, 29.33 km NW Manantenina, -24.13993 , 47.07418 , 540 m , montane rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); Anosy Region, PN Andohahela, Col de Tanatana, -24.7585 , 46.85367 , 275 m , rainforest (B.L. Fisher, F.A. Esteves et al.) (CAS); Foret Mandena 8.5 km N Tolagnaro, -24.95267 , 47.0025 , 20 m , littoral rainforest (B.L. Fisher et al.) (CAS); Isaka-Ivondro; Taolagnaro, -24.8, 46.866667, 41 m (Remy) (CAS); Ivohibe; Mahabo; Betroka, -23.62778 , 46.48861 , 1552 m (R. Decary) (CAS); Kirindy Forest, plot I, -20.07354 , 44.67194 , 60 m , dry forest (E. Lokensgard) (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); PN Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro, -24.76389 , 46.76683 , 650 m , rainforest (Fisher-Griswold Arthropod Team) (CAS); Mikea Forest, deciduous dry forest, -22.90367 , 43.4755 , 30 m , deciduous dry forest (R. Harin'Hala ) (CAS); Mikea Forest, spiny forest, -22.91333 , 43.48222 , 37 m , spiny forest (R. Harin'Hala ) (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). Diagnosis. With head in full-face view, lateral cephalic margins converge posteriorly towards eye level, strongly converging from posterior ocular margin to occipital corner of head; anteromedian margin of clypeus straight; two apical teeth of mandible normally spaced; lateral cephalic margin anterior to eye level covered with erect hairs. Description. Minor worker. In full-face view, lateral cephalic margins converging progressively towards posterior margin or slightly converging to level of anterior ocular margin and strongly converging posteriorly; eye large and convex (EL/CS: 0.28 +/- 0.02; 0.25-0.31), breaking lateral cephalic border, located at midlength of head, level of its posterior margin within the posterior 1/3 of head (PoOc/CL; 0.33 +/- 0.01; 0.31-0.36); frontal carinae more or less wide, posteriorly parallel (FR/CS: 0.25 +/- 0.01; 0.24-0.27), their distance larger than smallest distance to eye; clypeus with anterolateral angle and medially slightly concave anteromedian margin; two apical teeth of mandible distantly spaced; antennal scape relatively long (SL/CS: 1.93 +/- 0.08; 1.76-2.12). Mesosoma long and low (MPH/ML: 0.29 +/- 0.01; 0.27-0.32), with slightly convex promesonotum, nearly flat mesopropodeum, and feebly visible metanotal groove; junction of propodeal dorsum to declivity bluntly angulate; propodeal declivity 1/3 length of dorsum. Petiole nodiform, tapering dorsally; its dorsal margin inclined posteriorly and forming a blunt angle to anterior face; posterior face 3 x as high as the anterior; femur of hind leg rounded axially, not twisted basally. First and second gastral tergites without a pair of white spots; lateral margin of head covered with erect hairs; near posterior margin of head with more than six erect hairs; antennal scape covered with erect hairs inclined at ca. 45°; pronotum with numerous erect to suberect hairs; posterodorsal angle of propodeum with a pair of erect hairs. Major worker. With characteristics of minor worker except for the following divergent characters: larger head (CS: 3.93 +/- 0.51; 3.46-4.54; CWb/CL: 0.88 +/- 0.08; 0.80-0.97), with concavity on posterior margin; anteromedian margin of clypeus slightly concave; apical 1/3 of antennal scape surpassing posterior cephalic margin; promesonotum and propodeum in separate convexity; mesonotum and propodeal dorsum 2 x as long as declivity, with blunt angle junction. Distribution and biology. Camponotus dufouri is only known from Madagascar. Its distribution follows that of eastern humid forests ranging from the littoral to the montane rainforest of the island (Fig. 52G ). Its colony nests have been found mostly in rotten logs, rotten sticks, and rotting tree stumps, and rarely in dead branches, twigs, and bamboo on the ground. Workers forage frequently on the ground and in leaf litter, rarely on low vegetation. Figure 52. Camponotus dufouri A, C head in full-face view B, D lateral view E, F dorsal view of minor workers CASENT0060837 and CASENT0274127 G distribution map. Discussion. A clypeus with a straight anteromedian margin makes C. dufouri easy to separate from C. lokobe . Also, in C. tendryi , when the head is set in full-face view, the lateral cephalic margin posterior to the eye level is without erect hairs and its body size is smaller (CS: 1.57 +/- 0.01; 1.56-1.58), while in C. dufouri , erect hairs are present on the lateral cephalic margin posterior to the eye level and its body size is larger (CS: 1.97 +/- 0.19; 1.66-2.26). A verification of the syntype workers of C. dufouri imerinensis demonstrates that they were indistinguishable from the syntypes of C. dufouri . Moreover, since the geographic distribution of C. dufouri is ranging from the littoral to the montane rainforests of Madagascar where the type specimens of the subspecies name were collected, it is consequently correct to synonymize it under C. dufouri . The qualitative morphology-based study of C. dufouri agrees with the multivariate statistical analysis and both methods support the taxonomic delimitation for this species. The validity of the species, supported by the grouping obtained from NC-clustering, is confirmed by LDA with an identification success of 100%.