Tetraponera grandidieri

Ward, P. S., 2009, The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae)., Journal of Hymenoptera Research 18, pp. 285-304 : 288-294

Tetraponera grandidieri (Forel 1891)

(Figs 7-8, 19)

Sima Grandidieri Forel 1891: 203. Syntypes, 4 workers, Central Madagascar (Hildebrandt) ( MCSN, MHNG) [examined] [Two of three MHNG syntypes imaged on AntWeb: CASENT0101652 , CASENT0102029 ]. One syntype ( CASENT0101652 ) here designated lectotype .

Figs 1-6. Workers of the Tetraponera grandidieri group, right mandible (1, 2), posterior view of propodeum (3, 4), and lateral view of mesosoma (5, 6). 1, T. merita ; 2, T. hespera ; 3, T. hespera, Ankarana population; 4, T. hespera, Nosy Be; 5, T. inermis ; 6, T. hespera .

Sima Grandidieri var. Hildebrandti Forel 1891: 204. Holotype (by monotypy) worker, Pays de Betsileo, "Sud Central Madagascar" (Hildebrandt) (MHNG) [examined] [Imaged on AntWeb: CASENT0101883]. Syn. n.

Sima grandidieri Forel ; Forel 1891: 229-230. Description of queen and male.

Tetraponera grandidieri (Forel) ; Wheeler 1922: 1014. Combination in Tetraponera .

Tetraponera grandidieri var. hildebrandti (Forel) ; Wheeler 1922: 1014. Combination in Tetraponera .

Tetraponera grandidieri (Forel) ; Ward, 1991: 342. Nesting biology.

Tetraponera grandidieri (Forel) ; Fisher 1996: 100; Fisher 1999: 134; Fisher 2002: 318. Cited in faunal inventories.

Tetraponera cf. grandidieri (Forel) ; Fisher 1998: 49. Cited in faunal inventory.

Tetraponera grandidieri (Forel) ; Ward and Downie 2005. DNA sequences of five nuclear genes; GenBank accession numbers AY703507 (18S rDNA), AY703574 (28S rDNA), AY703641 (wingless), AY703775 (long wavelength rhodopsin), and AY703778 (abdominal-A).

Material Examined.-(BMNH, CASC, MCSN, MCZC, MHNG, MNHN, NHMB, NHMV, PSWC, SAMC, UCDC, USNM) MADAGASCAR Antananarivo: Andrangoloaka (Sikora); Antsiranana: Montagne d'Ambre [?] [as "Amber geb."] (Rolle); 1 km W Andampibe, Cap Masoala, 125 m (Alpert, G. D.); 3 km W Andampibe, Cap Masoala, 125 m (Alpert, G. D.); 3 km W Sakalava Beach, 40 m (Schlinger; et al.); 4 km SW Ambohitra (=Joffreville), 1000 m (Ward, P. S.); 5 km SW Ambohitra (=Joffreville), 1100 m (Ward, P. S.); 7 km N Joffreville, 360 m (Harin'Hala, R.); Betaolana forest, 880 m (Fisher, В. L.; et al.); Diego Suarez (Alluaud, C); Forêt Ambanitaza, 26.1 km 347° [NNW] Antalaha, 240 m (Fisher, B. L.; et al.); Forêt Binara, 9.1 km 233° SW Dairana, 650-800 m (Fisher, B. L.; et al.); Forêt Binara, 9.4 km 235° SW Dairana,' 1100 m (Fisher, B. L.; et al.); Fotodriana, Cap Masoala, 25 m (Alpert, G. D.; et al.); Fotodriana, Cap Masoala, 25 m (Alpert, G. D.); Marojejy R.N. I., #12, 375 m (Alpert, G. D.); Marojejy R.N. I., #12, 665 m (DiRosa, R.); Montagne d'Ambre, 900 m (Alpert, G.; et al.); Montagne d'Ambre, Petit Lac, 1000 m (Alpert, G.; et al.); Montagne Français, 150 m (Harin'Hala, R.); P.N. Marojejy, 26.6 km 31° NE Andapa, 1325 m (Fisher, B. L.; et al.); P.N. Marojejy, 27.6 km 35° NE Andapa, 775 m (Fisher, B. L.; et al.); P.N. Marojejy, 28.0 km 38° NE Andapa, 450 m (Fisher, B. L.; et al); P.N. Montagne Ambre, 1125 m (Harin'Hala, R.); P.N. Montagne Ambre, 3.6 km 235° SW Joffreville, 925 m (Fisher, B. L.; et al.); P.N. Montagne Ambre, 960 m (Harin'Hala, R.); P.N. Montagne Ambre, 960 m (Irwin, M. E.; et al.); P.N. Montagne Ambre, 960 m (Schlinger; et al.); P.N. Montagne Ambre, 975 m (Irwin, M. E.; et al.); Pare Nat. Mont. d'Ambre [as "Amber Mt. Nat. Pk."], 3000 ft. (Alpert, G. D.); Pare Nat. Mont. d'Ambre [as "Amber Mt. Nat. Pk."], 3200 ft. (Alpert, G. D.); Pare Nat. Mont. d'Ambre [as "Amber Mt. Nat. Pk."], 3600 ft. (Alpert, G. D.); Pare Nat. Mont. d'Ambre, 1000-1100 m'(Brown, W. L.; Brown, D. E.); Pare Nat. Montagne d'Ambre, 1100 m (Oison, D. M.); R.S. Manongarivo, 10.8 km 229° SW Antanambao, 400 m (Fisher, В. L.); R.S. Manongarivo, 12.8 km 228° SW Antanambao, 780 m (Fisher, В. L.); R.S. Manongarivo, 14.5 km 220° SW Antanambao, 1175 m (Fisher, В. L.); Réserve Spéciale Ambre, 3.5 km 235° SW Sakaramy, 325 m (Fisher, В. L.; et al.); Sakalava Beach, 10 m (Harin'Hala, R.); Fianarantsoa: 3 km W Ranomafana, nr. Ifanadiana, 950 m (Ward, P. S.); 40 km S Ambalavao, Rés. Andringitra, 1275 m (Fisher, B. L.); 45 km S Ambalavao, 720 m (Fisher, B. L.); 7 km W Ranomafana Nati. Park, 1000 m (Steiner, K.); 7 km W Ranomafana Nati. Park, 1000 m (Steiner, W. E.; Zack, S.); 7 km W Ranomafana, 1000 m (Stebbins, M.; et al); 7 km W Ranomafana, 900 m (Steiner, W. E.); 8 km E Kianjavato, 145 m (Alpert, G.); 9 km ESE Ranomafana, nr. Ifanadiana, 600 m (Ward, P. S.); FC Vatovavy, 175 m (Fisher, B. L.; et al.); Ivohibe, 1500 m (Decary, R.); Maharira Forest, Ranomafana Nati Pk, 1350 m (Rajeriarison, E.); Maharira Forest, Ranomafana Nati Pk, 1375 m (Rajeriarison, E.); Miaranony, Ranomafana Nati Pk, 1050 m (Rajeriarison, E.); Miaranony, Ranomafana Nati Pk, 700 m (Rajeriarison, E.); Pays de Betsileo, "Sud Central Madagascar" (Hildebrandt); P.N. Ranomafana, 0.4 km WSW park entrance, 900 m (Kavanaugh, D. H.; Kavanaugh, K. M.); P.N. Ranomafana, 1020 m (Harin'Hala, R.); P.N. Ranomafana, 1130 m (Harin'Hala, R.); P.N. Ranomafana, Vatoharanana, 4.1 km 231° SW Ranomafana, 1100 m (Fisher, B. L.; et al.); P.N. Ranomafana, Vohiparara, 1110 m (Harin'Hala, R.); PN Befotaka-Midongy, 940 m (Fisher, B. L.; et al.); R.S. Ivohibe, 8.0 km E Ivohibe, 1200 m (Fisher, B. L.); Ranomafana (Pauly, A.); Ranomafana N. P., 1000 m (Alpert, G.; et al.); Ranomafana N.P., Talatakely Forest, Piste S 100, 900 m (Rajeriarison, E.); Ranomafana N.P., Vohiparara Forest, 1200 m (Rajeriarison, E.); Ranomafana National Park, Talatakely, 915- 1000 m (Lee, V. F.; Ribrado, K. J.); Ranomafana Nati Pk. (Rajeriarison, E.); Ranomafana Nati Pk., 950-1100 m (Bartolozzi, L.; et al.); Ranomafana Nati. Pk., Saharoemba ZP, 800 m (Rabeson, P.); Ranomafana NP, Talatakely (Griswold, C. E.; Ubick, D.); Ranomafana, Ambatolahy forest (Rajeriarison, E.); Ranomafana, Ambatovory forest, 1035 m (Rajeriarison, E.); Ranomafana, Miaranony Village (Kingman, A.); Ranomafana, Vohiparara forest, 1160 m (Rajeriarison, E.); Vevembe, 600 m (Fisher, В. L.; et al.); Toamasina: 10 km S Cap Este, 5 km W, 20 m (Alpert, G. D.); 14 km W Cap Est, Ambato, 100 m (Alpert, G. D.); 17 km W Andapa, Rés. d'Anjanaharibe-Sud, 875 m (Alpert, G. D.); 19 km ESE Maroantsetra, 250 m (Ward, P. S.); 19 km ESE Maroantsetra, 300 m (Ward, P. S.); 19 km ESE Maroantsetra, 350 m (Ward, P. S.); 1 km SSW Andasibe (= Périnet), 920 m (Ward, P. S.); 6.3 km S Ambanizana, Andranobe, 25 m (Fisher, В. L.); 6.5 km SSW Befingotra, Res. Anjanaharibe-Sud, 875 m (Fisher, В. L.); 6.9 km NE Ambanizana, 1080 m (Fisher, В. L.); 6.9 km NE Ambanizana, 650 m (Fisher, В. L.); 8 km ESE Andasibe (= Périnet), 800 m (Ward, P. S.); Ambodiriana, 125 m (Fisher, В. L.; et al.); Ampasimbe, 450 m (Betsch, J.-M.); Andasibe ( Périnet) (Brooks, R. W.); Andasibe PN, 1025 m (Harin'Hala, R.); Anosibe An'ala [as " Nosibé, Village de l'Imerina "] (Sikora); Antongil (Mocquerys); Asanoreyo, 3 km W Anosibe An'ala (Raharimina, C); Baie d'Antongil (eu.); Betampona, 520 m (Fisher, В. L.; et al.); Kalalao, 100 m (Fisher, B. L.; et al.); Manakambahiny (Pauly, A.); Mont. Akirindro, 7.6 km 341° NNW Ambinanitelo, 600 m (Fisher, В. L.; et al); Mont. Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 470 m (Fisher, В. L.; et al.); Mont. Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 1100 m (Fisher, В. L.; et al.); Nosey Mangabe (Alpert, G. D.); Nosy Mangabe, <5 m (Ward, P. S.); Nosy Mangabe, 150 m (Ward, P. S.); Nosy Mangabe, 20 m (Ward, P. S.); Nosy Mangabe, 300 m (Ward, P. S.); Nosy Mangabe, 3 m (Fisher, В. L.; et al.); P.N. Mantadia, 895 m (Ratsirarson, H. J.); Permet ( Noyés, J. S.; Day, M. C); Permet (Ross, E. S.); PN Mananara-Nord, 225 m (Fisher, В. L.; et al.); PN Zahamena, 860 m (Fisher, В. L.; et al.); PN Zahamena, Besaky River, 760 m (Fisher, В. L.; et al.); PN Zahamena, Oribe River, 780 m (Fisher, В. L.; et al.); PN Zahamena, Sahavorondrano River, 765 m (Fisher, В. L.; et al.); Saint Marie, Forêt de Kalalao (Madl); St. Marie (eu.); Tampolo, 218 m (Fisher, В. L.; et al.); Toliara: 10 km NW Enakara, Rés. Andohahela, 420 m (Fisher, В. L.); 10 km NW Enakara, Rés. Andohahela, 430 m (Fisher, В. L.); 10 km SSW Eminiminy, 750 m (Rajeriarison, E.); 11 km NW Enakara, Rés. Andohahela, 800 m (Fisher, В. L.); 5 km NNW Isaka-Ivondro, Rés. Andohahela, 280 m (Ward, P. S.); 5 km WNW Mandiso, Rés. Andohahela, 400 m (Ward, P. S.); Col de Manangotry, c.30 km N Fort Dauphin, -1000 m (Whitacre, D.); Forêt Ivohibe, 650 m (Fisher, В. L.; et al); Fort Dauphin (Alluaud, C); Manatantely, 100 m (Fisher, В. L.; et al.); P.N. Andohahela, 3.8 km 113° ESE Mahamavo, 900 m (Fisher, В. L.; et al.); P.N. Andohahela, Manampanihy,5.4 km 113° ESE Mahamavo, 650 m (Fisher, В. L.; et al); PN Andohahela, 275 m (Fisher, B. L.; et al.); Rés. Andohahela, Marosohy, 600 m (Fisher, В. L.); Vallee d'Ambolo, Col de Sakalavana (Alluaud, C); province unknown: "Central Madagascar" (Hildebrandt); "Centre de Madag" (Hildebrandt); "Madag." (Sikora); "Madagascar Centralis" (Hildebrandt); "Madagascar" (cu.); "Madagascar" (de Gaulle, J.); "Madagascar" (Grandidier); "Madagascar" (Sikora); "Madagascar/(S.-E.)" (Decary, R.).

Worker measurements (n = 13). HW

1.01-1.44, HL 1.20-1.65, LHT 1.07-1.56, CI 0.77-0.88, FCI 0.15-0.17, REL 0.28-0.36, REL2 0.34-0.43, SI 0.74-0.81, FI 0.29-0.36, PLI 0.50-0.59, PWI 0.40-0.53.

Worker diagnosis. With characteristics of the T. grandidieri group (see above); basal margin of mandible edentate; anterior clypeal margin broadly convex and crenulate, directed forward, not anteroventrally; head relatively elongate (CI 0.77-0.88); metanotal spiracle more or less visible in lateral view of mesosoma, protruding dorsally in the mesopropodeal impression; dorsal face of propodeum broadly convex in lateral and posterior views; standing pilosity generally sparse; long paired setae (0.2-0.4 mm in length) distributed as follows: 1 pair between the frontal carinae, 1 pair on upper half of head, 1 pair on the pronotum, 0-2 pairs on the petiole; 1-2 pairs on the postpetiole; standing pilosity scattered on successive abdominal segments(gastric segments 1-4); short appressed to subdecumbent hairs absent or inconspicuous on most of body; integument mostly sublucid, with fine coriarious /puncticulate sculpture; body orange-brown, appendages lighter; head usually dark brown to brownish-black, but concolorous with rest of body in some northern populations (see discussion below); legs uniformly light orange-brown.

Comments. This species is typically bicolored with a black or dark brown head and the remainder of the body a contrasting orange-brown. This allows it to be distinguished from the other two species, Г. inermis and Г. merita , with which it is widely sympatric -both of these usually have the head more or less concolorous with the mesosoma. Some northern populations of T. grandidieri have workers that are unicolorous orange-brown, however, and these superficially resemble the other two species. They can be recognized because they lack a tooth on the basal margin of the mandible ( présent in T. merita ) and the metanotal spiracle protrudes from the mesosoma dorsum in profile (not protruding in T. inermis ). The degree of prominence of the metanotal spiracle varies, however, so it is also useful to examine head shape, which is more elongate in T. grandidieri (worker CI 0.77- 0.88 versus 0.88-0.97 in T. inermis ; see also additional discussion under T. inermis ). T. grandidieri also overlaps in distribution with T. hespera in northern Madagascar. Where these two species co-occur T. grandidieri has a bicolored body, while T. hespera has a unicolored body and contrasting dark bands on the femora.

At Betampona (17°53'S 49°12'E) Brian Fisher collected three nest séries of T. grandidieri : one (BLF13292) with unicolored workers, a second (BLF13298) with bicolored workers, and a third (BLF13349) with both unicolored and bicolored workers, in approximately equal proportions. The Betampona workers with light and dark heads show no obvious différences other than color. The occurrence of both forms in the same nest is consistent with the view that they are conspecific. In addition, genetic data (>10 kb of sé- quence data from several nuclear genes and one mitochondrial gene) from populations sampled throughout the range of the species show the two color forms to be phylogenetically comingled (Ward unpubl.).

Both color forms are here treated as conspecific but further studies are needed to clarify their status. It is possible that these color morphs show some degree of reproductive isolation and/or ecotypic differentiation. As indicated below, they appear to be involved in a mimicry complex with some species of Camponotus .

Finally it should be noted that there are nine specimens of T. grandidieri in the Forel collection in MHNG (Geneva) labeled as "Typus" or "Cotypus" but most are not true types, because the label data exclude this possibility. These non-types include three males (from Andrangoloaka), one dealate queen (from Andrangoloaka) and one worker (from " Nosibé, Village de lTmerina"), ali with a red "Typus" label, and an alate queen (Madagascar/Sikora) labeled "Cotypus". Only three workers in MHNG are apparently part of the actual type séries of Г. grandidieri (there is also a syntype worker in MCSN). To avoid confusion I have designated one of the MHNG syntype workers as lectotype.

Distribution and biology. Tetraponera grandidieri is widespread in eastern Madagascar, with a distribution that spans the length of the island (Fig. 23). Populations are restricted to rainforest, at élévations ranging from sea level to 1375 m. As a result of habitat destruction in the lowlands most populations are found at intermediate or higher élévations. Colonies usually occupy dead twigs or branches on the ground, less commonly in the lower canopy. During field work in Madagascar I collected thirteen nest séries of this species, of which nine were in dead wood and four were located in cavities of live plants: three in stems of tree saplings (Ixora sp., Leea sp. and an unidentified plant), and one in a cavity in a live root of a tree in the genus Rhus. There were no scale insects (Coccoidea) in any of these live cavity nests, however, and there is no indication that T. grandidieri is closely associated with any particular plant species. It seems clear that it and other members of the T. grandidieri group occupy moister nest sites than most Tetraponera species. The nests of T. grandidieri apparently contain no more than one dealate queen, and colony sizes are small (5-40 workers). Alate queens and males have been collected from February to May. Workers commonly forage on low végéta- tion, and they appear to be mimicked by members of the Camponotus putatus complex whose workers forage in similar microhabitats. T. grandidieri is generally absent from disturbed rainforest edge and other high light environments.