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The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae) PDF

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Preview The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae)

HYM. RES. J. Vol. 18(2), 2009, pp. 285-304 The Ant Genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae) Ward Philip S. Department of Entomology, University of California, Davis, CA 95616, USA — Abstract. Ants in the Tetraponera grandidieri group are endemic to the island of Madagascar, where they occur in relatively undisturbed mesic forest. In this taxonomic revision of the group sevenspeciesarerecognized: T.grandidieri(Forel), T. hespera sp.n., T. hirsutasp.n., T. inermis sp.n., T. manangotrasp.n., T. meritasp.n. andT. variegata (Forel) stat.n. T.grandidierihildebrandti(Forel)is proposedasanewsynonymofT.grandidieri. Thespeciesinthis group showlimitedmorphological and genetic divergence. The justification for treating them as different species is that they occur sympatrically in various combinations, without showing genetic or phenotypic intergradation. Although differences in shape, pilosity and sculpture are not pronounced, there is notable color pattern variation, both within and among species. The conspicuous orange and reddish-brown colorthatcharacterizestheworkersandqueenslikelyservesaswarningcoloration.Theseantshave painful stings and several species ofants in the Camponotus putatus complex exhibit color patterns that apparently mimic those of the T. grandidieri group. Tw^ig-dwelling ants in the subfamily biological species concept (Mayr 1963; Pseudomyrmecinae are a distinctive com- Coyne and Orr 2004). ponent ofthe arboreal ant fauna in forests Thispaperis dedicatedtothememoryof and woodlands of both the Neotropics Roy Snelling, a colleague, friend and and Paleotropics (Ward and Downie ardent hymenopterist. In his later years 2005). The Afrotropical representatives of Roy developed an interest in the ant fauna the subfamily, currently placed in the of the Afrotropical region, specifically that genus Tetraponera F. Smith, were recently of Kenya, and his last days were spent divided into five monophyletic species there. Roy's generosity, candor, pungent groups (Ward 2006). Four of the five humor, and enthusiasm for ants and other groups occur in Madagascar and one of aculeate Hymenoptera left an indelible these, the Tetraponera grandidieri group, is impression on those who had the pleasure endemic to the island. The group has of interacting with him. never received the benefit of a modern MATERIALS AND METHODS taxonomic treatment. There is only a single named species, T. grandidieri Specimens were examined in the follow- (Forel), with two nominal subspecies, but ing collections: the current study reveals substantially greater species-level diversity, paralleling BMNH Natural History Museum, UK the situation for the ant fauna of Mada- London, gascar as a whole, where considerable CASC California Academy of Scienc- numbers of species remain undescribed es, San Francisco, CA, USA (Fisher 2003). Species are here delimited CUIC Cornell University Insect Col- using a combination of morphological, lection, Ithaca, NY, USA geographical and genetic evidence, while MCSN MuseoCivicodeHistoriaNatural working within the framework of the "Giacomo Doria'', Genoa, Italy 286 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling MCZC Museum of Comparative Zool- FW Maximum width of profemur, ogy, Harvard University, Cam- measured in the same view as bridge, MA, USA FL and at right angles to it. MHNG Museum d'Histoire Naturelle, PL Length of the petiole in lateral Geneva, Switzerland view from the lateral flanges of MNHN Museum National d'Histoire the anterior peduncle to the Naturelle, Paris, France posterior margin of the petiole. NHMB Naturhistorisches Museum, PH Maximum height of petiole, Basel, Switzerland measured in the same view as NHMV Naturhistorisches Museum, PL, and excluding protruding Vienna, Austria teeth or lobes at the anteroven- PSWC P. S. Ward Collection, Univer- tral or posteroventral extremi- sity of California at Davis, CA, ties of the petiole. USA DPW Maximum width of petiole, SAMC South African Museum, Cape measured in dorsal view. Town, South Africa LHT Length of the metatibia, ex- UCDC Bohart Museum of Entomolo- cluding the proximomedial gy, University of California at condyle (Ward 2001, fig. 5). HW/HL Davis, CA, USA CI Cephalic index: USNM National Museum of Natural FCI Frontal carinaindex: MFC/HW History, Washington, DC, USA REL Relative eye length: EL/HL REL2 Relative eye length, using HW: Standard measurements (in mm) and EL/HW setal counts were taken at 50X with a Wild SI Scape index: SL/HW MSA microscope, as described in Ward FI Profemur index: FW/FL (2001, 2006). The abbreviations used for PLI Petiole length index: PH/PL measurements, indices and setal counts are PWI Petiole width index: DPW/PL given below. The first four measurements CSC Cepalic setal count: number of are taken with the head in full-face view, standing hairs (those forming such that the posterior margin of the head an angle of 45° or more with and the anterolateral corners are in the the cuticular surface) visible on same plane of view. the posterior half of the head, HW Maximum head width, includ- as seen in lateral and posterior views HL Hinegaedyelse.ngth, taken along the MSC Mesosomal setal count: num- ber of standing hairs visible in midline, from the posterior profile (lateral view) on the margin of the head to the mesosoma dorsum anterior extremity of the clyp- eus. Automontage images of selected speci- EL Eye length, measured in the mens (Figs 7-22) were taken by April same plane of view as HL. Nobile and Erin Prado at the California MFC Minimum distancebetweenthe Academy of Sciences (CAS), under the frontal carinae. direction of Brian Fisher. These images SL Scape length, excluding the are also posted on AntWeb (www.antweb. radicle. org), togetherwithphotographs ofthe type FL Length of profemur, measured specimens of T. grandidieri, T. grandidieri along its long axis in posterior hildebrandti (Forel) and T. grandidieri var- view. iegata (Forel). Volume 18, Number1, 2009 287 The species described here were se- with four teeth; basal margin with 0-1 quenced for fragments of one mitochon- teeth and subequal inlength to masticatory drial gene (COI) and several nuclear genes, margin; labrum with a pair of tubercles using methods described in Ward and closely flanking the midline near the Downie (2005) and Brady et al. (2006). This proximal margin but lacking a median molecular work is ongoing and results will tubercle; palp formula 6,4; anteromedial be analyzed and presented in more detail margin of clypeus crenulate or emarginate; DNA elsewhere. The sequence data pro- distance betw^een frontal carinae exceeding vide ancillary information that helps to basal scape width (FCI 0.11-0.18), scape validate species boundaries inferred from length three-quarters or more of head morphology and geography. width (SI 0.72-0.83); eye length about Species distributions were plotted with one-third of head length (REL 0.28-0.36); the shareware program Versamap (Version head capsule with three distinct ocelli; 3.01). For most specimens examined in this pronotum laterally marginate, but not study the coordinates (latitude and longi- strongly so; mesopropodeal impression tude) were given on the specimen label. well developed (Figs 5, 6); petiole relative- For material lacking this information the ly long (PLI 0.49-0.59, PWI 0.40-0.65); following sources were used to georefer- posteroventral margin of petiole lying ence collection sites: Forel (1892), United adjacent to helcium venter; metabasitarsal States Board on Geographic Names (1989), sulcus present; legs long and slender (FI Viette (1991), Ruber (2003), the GEOnet 0.28-0.36, LHT/HL 0.85-1.12); appressed Names Ser\^er (http://earth-info.nga.mil/ pubescence sparse on abdominal tergite 4; gns/html/index.html), the Gazetteer to standing pilosity uncommon (CSC 2-3, Malagasy Botanical Collecting Localities MSC 1-6), absent from mesonotum, propo- (http://ww^w.mobot.org/MOBOT/Research/ deum, and extensor surfaces of the tibiae. madagascar/gazetteer/), and topographic Orange to reddish-brown, head concolor- maps of Madagascar at scales of 1:50,000, ous or darker; gaster and portions of 1:100,000 and 1:500,000, published by Foi- femora may also be infuscated. ben-Taosarintanin' i Madagasikara (Insti- Comments. Distinctive features of the tut Geographique et Hydrographique Na- worker caste of the T. grandidieri group tional). In the lists of material examined, include the relatively large body size, long most locality names are given verbatim legs and antennal scapes, presence of three from the specimen label, but in a few oceUi, deeply impressed mesopropodeal instances they have been interpreted for impression, and conspicuous orange to clarity. In this case the original spelling is reddish-brown body coloration. Other given in quotes after the emendation (e.g., Malagasy Tetraponera species have shorter Anosibe An'ala [as "Nosibe, Village de scapes and legs (SI 0.40-0.70, LHT/HL ITmerina"]). The abbreviation ''c.u." signi- 0.58-0.82), 0-2 ocelH on the head, a fies collector unknown. shallower mesopropodeal impression, and usually darker body color. Additional RESULTS differences between the T. grandidieri group and the other four species groups Diagnosis of the Tetraponera grandidieri of Afrotropical Tetraponera are given in group Ward (2006). (modified from Ward 2006) Synonymic list of species Worker diagnosis. Medium to large r. grandidieri (Forel 1891: 203) species (FIW 0.95-1.59, HL 1.05-2.01, LHT = T. grandidieri hildebrandti (Forel 1891: 203) 1.05-1.83); masticatory margin of mandible syn. n. 288 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling T. hespera sp. n. T. manangotra sp. n. T, hirsuta sp. n. T. merita sp. n. T. inennis sp. n. T. variegata (Forel 1895: 487) stat. n. KEY TO SPECIES BASED ON THE WORKER CASTE 1 Basal margin of mandible with a prominent tooth, in addition to four teeth on the masticatory margin (Fig. 1); anterior clypeal margin deflected ventrally; wide- spread in eastern and northern Madagascar merita - Basal margin of mandible lacking tooth, masticatory margin with four teeth (Fig. 2); anterior clypeal margin directed forward, not deflected ventrally 2 2(1) Petiolebroad (PWI 0.61-0.65, DPW/HW 0.50-0.53), subtriangularin dorsal view, and HW witharelativelyshort,thickanteriorpeduncle(Fig. 20);largerspecies, 1.48-1.58, LHT 1.64—1.76; knownonlyfrom extreme southern Madagascar manangotra Petiole more slender (PWI 0.40-0.53, DPW/HW 0.30-0.40), obovate, and with a thin, HW elongate anterior peduncle (e.g.. Figs 14, 16, 18); smaller species, 0.95-1.44, LHT 1.05-1.59; widespread 3 3(2) Scape with conspicuous suberect and subdecumbent hairs (Fig. 13); body tricolored: metasoma, appendages, and ventral margin of mesosoma orange, most of mesosoma reddish-brown,andheaddarkbrownishblack;endemictoManongarivoMassif hirsuta - Most hairs on scape appressed or decumbent, and generally inconspicuous, except those at the apex (e.g.. Fig. 9); body color variable but usually without preceding tricolor pattern 4 4(3) Metanotalspiraclenotprotrudingabovetheprofileofthemesosoma,asseeninlateral view (Fig. 5); head broad (CI 0.88-0.97); head and mesosoma reddish-brown, metasoma and appendages paler; widespread in eastern Madagascar inermis - Metanotalspiraclemoreorlessprotrudingabovetheprofileofthemesosoma,asseenin lateral view (Fig. 6);head usuallymore elongate (CI 0.77-0.90); colorvariable .... 5 5(4) Dorsum of propodeum laterally compressed, the propodeum appearing subtriangu- lar in posterior view (Fig. 3); body concolorous orange-brown; northern Mada- gascar hespera, in part (Ankarana population) - Dorsum of propodeum more broadly rounded, the propodeum appearing dome- shaped in posterior view (Fig. 4); color variable 6 6(5) Legs uniformly light orange-brown, femora lacking conspicuousblackbanding;body usually bicolored, such that dark head contrasts with lighter orange-brown mesosoma and metasoma (Fig. 8), less commonly unicolorous orange; widespread and variable species grandidieri - Legs light orange-brown, with contrasting black bands on the distal portions of the mesofemur and metafemur (Figs 10, 22); body concolorous or bicolored (in latter case both head and gaster are dark brownish-black) 7 7(6) Body concolorous yellow brown or orange brown (Fig. 10); northwestern Madagascar hespera, in part (most populations) - Body bicolored, head and gaster dark brown and contrasting with the lighter mesosoma (Fig. 22); eastern Madagascar variegata Tetraponera grandidieri (Forel 1891) (MCSN, MHNG) [examined] [Two of three MHNG (Figs 7-8, 19) syntypes imaged on AntWeb: CASENT0101652, CASENT0102029]. One Sima Grandidieri Forel 1891: 203. Syntypes, 4 syntype (CASENT0101652) here designated workers. Central Madagascar (Hildebrandt) lectotype. . VoLL-ME 18, Xl-mber2, 2009 289 0.5mm I 0.5 mm 1 mm 0.5 Figs 1-6. Workersofthe Tetraiponeragrandidierigroup,rightmandible (1,2),posteriorvie^v ofpropodeum (3, 4),andlateralviewofmesosoma(5,6). 1,T. nierita;2,T.Jiespera;3,T.hespera,Ankaranapopulation;4,T. hespera, NosyBe; 5, T. inemiis; 6, T. hespera. Sima Grandidieri var. Hildebrandti Forel 1891: Material Examined.—{V>MNH, CASC, MCSN, 204. Holot}^e (by monoty^py) worker. Pays MCZC, MHNG, MNHN, NHMB, NHMV, de Betsileo, "Sud Central Madagascar" (Hil- PSWC, SAMC, UCDC, USNM) MADAGAS- debrandt) (MHNG) [exarriined] [Imaged on CAR Antananarivo: Andrangoloaka (Sikora); AntWeb: CASEXT0101883]. Syn. n. Antsiranana: Montague d'WAmbre [?] [as "Amber Sima grandidieri Forel; Forel 1891: 229-230. geb."] (Rolle); 1 km Andampibe, Cap Description of queen and male. Masoala, 125 m (Alpert, G. D.); 3 km W m Tetraponera grandidieri (Forel); WH-ieeler 1922: Andampibe, Cap Masoala, 125 (Alpert, G. 1014. Combination in Tetraponera. D.); 3 kmW Sakalava Beach, 40 m (Schlinger; et Tetraponera grandidieri var. hildebrandti (Forel); al.); 4 km SW Ambohitra (=Joffreville), 1000 m Wheeler 1922: 1014. Combination in Tetra- (Ward,P. S.);5kmSWAmbohitra(=Joffreville), ponera. 1100 m (Ward, P. S.); 7 km N Joffreville, 360 m m Tetraponera grandidieri (Forel); Ward, 1991: 342. (Harin'Fiala,R.);Betaolanaforest,880 (Fisher, Nesting biology. B. L.; et al.); Diego Suarez (Alluaud, C); Foret Tetraponera grandidieri (Forel); Fisher 1996: 100; Ambanitaza,26.1 km347^ [NNW] Antalaha,240 Fisher 1999: 134; Fisher 2002: 318. Cited in m (Fisher, B. L.; etal.);ForetBinara,9.1 km233= faunal inventories. SW Dairana, 650-800 m (Fisher, B. L.; et al.); Tetraponera cf.grandidieri (Forel);Fisher 1998: 49. Foret Binara, 9.4 km 235^ SW Dairana, 1100 m Cited infaimal inventor}' (Fisher, B. L.; et al.); Fotodriana, Cap Masoala, Tetraponera grandidieri (Forel); W^ard and 25 m (Alpert, G. D.; et al.); Fotodriana, Cap Downie 2005. DNA sequences offive nuclear Masoala, 25 m (Alpert, G. D.); Marojejy R.N.I., m genes; GenBank accession numbers #12,375 (Alpert,G. D.);MarojejyR.N.I., =12, m m AY703507 (IBS rDNA), AY703574 (28S 665 (DiRosa, R.); Montague d'Ambre, 900 rDNA), AY703641 (wingless), AY703775 (Alpert, G.; et al.); Montague d'Ambre, Petit m (long wa^-elength rhodopsin), and AY703778 Lac, 1000 (Alpert, G.; et al.); Montague m (abdominal-A). Fraucais, 150 (Fiarin'Hala, R.); P.N. Marojejy, 290 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling m r^ 11 1 ) 'r^ \-?!5^y.\ ^: mm 0.5 Figs 7-12. AutomontageimagesofworkersoftheTetraponeragrandidierigroup,full-face(dorsal)viewofhead (7, 9, 11) and lateral view ofbody (8, 10, 12). 7, 8, T. grandidieri (CASENT0012861); 9, 10, T. hespera, holotype (CASENT0012865); 11, 12, T. hespera, Ankarana population (CASENT0012864). 26.6km ?>V NE Andapa, 1325 m (Fisher,B. L.;et Ambre, 960 m (Harin'Hala, R.); P.N. Montagne al.); P.N. Marojejy, 27.6 km 35^ NE Andapa, 775 Ambre, 960 m (Irwin, M. E.; et al.); P.N. m (Fisher, B. L.; et al); P.N. Marojejy, 28.0 km Montagne Ambre, 960 m (Schlinger; et al.); 38' NE Andapa,450m (Fisher,B. L.;etal.);P.N. P.N. Montagne Ambre, 975 m (Invin, M. E.; et MontagneAmbre, 1125 m (Harin'Hala, R.);P.N. al.); Pare Nat. Mont. d'Ambre [as "Amber Mt. Montagne Ambre, 3.6 km 235= SW Joffreville, Nat. Pk."], 3000 ft. (Alpert, G. D.); Pare Nat. m 925 (Fisher, B. L.; et al.); P.N. Montagne Mont. d'Ambre [as "AmberMt. Nat. Pk."],3200 Volume 18, Number1, 2009 291 0.5mm 2mm Figs 13-18. Automontage images of workers of the Tetraponera grandidieri group, full-face (dorsal) view of head (13, 15, 17) and lateral view ofbody (14, 16, 18). 13, 14, T. hirsuta, holot\^e (CASENTOl70370); 15, 16, T. inermis,holotype (CASENT0012862); 17, 18, T. merita, holotype (CASENT0012863). ft. (Alpert, G. D.); Pare Nat. Mont. d'Ambre [as SW Antanambao, 780 m (Fisher, B. L.); R.S. "Amber Mt. Nat. Pk."], 3600 ft. (Alpert, G. D.); Manongarivo, 14.5 km 220' SW Antanambao, m m PareNat. Mont. d'Ambre, 1000-1100 (Brown, 1175 (Fisher, B. L.); Reserve Speciale Ambre, W. L.; Brown, D. E.); Pare Nat. Montagne 3.5 km 235= SW Sakaramy, 325 m (Fisher, B. L.; m m d'Ambre, 1100 (Olson, D. M.); R.S. Manon- et al.); Sakalava Beach, 10 (Harin'Hala, R.); garivo, 10.8 km 229= SW Antanambao, 400 m Fianarantsoa: 3 km W Ranomafana, nr. Ifanadi- (Fisher, B. L.); R.S. Manongarivo, 12.8 km 228= ana, 950 m (Ward, P. S.); 40 km S Ambalavao, 292 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling ^^H 19 20 mm 0.5 Figs 19-22. Automontage images of workers of the Tetraponera grandidieri group, full-face (dorsal) view of head (19, 21) and lateral view ofbody (20, 22). 19, 20, T. manangotra, paratype (CASENT0121948); 21, 22, T. variegata (CASENT0409559). Res. Andringitra, 1275 m (Fisher, B. L.);45kmS Ranomafana, Vatoharanana, 4.1 km 231° SW Ambalavao, 720 m (Fisher, B. L.); 7 km W Ranomafana, 1100 m (Fisher, B. L.; et al.); P.N. Ranomafana Natl. Park, 1000 m (Steiner, K.); 7 Ranomafana, Vohiparara, 1110 m (Harin'Hala, km W Ranomafana Natl. Park, 1000 m (Steiner, R.); PN Befotaka-Midongy, 940 m (Fisher, B. L.; W. E.; Zack, S.); 7 km W RWanomafana, 1000 m et al.); R.S. Ivohibe, 8.0 km E Ivohibe, 1200 m (Stebbins, M.; et al); 7 km Ranomafana, 900 (Fisher, B. L.); Ranomafana (Pauly, A.); Rano- m (Steiner, W. E.); 8 km E Kianjavato, 145 m mafana N. P., 1000 m (Alpert, G.; et al.); (Alpert, G.); 9 km ESE Ranomafana, nr. Ifanadi- Ranomafana N.P., Talatakely Forest, Piste S ana, 600 m (Ward, P. S.); EC Vatovavy, 175 m 100, 900 m (Rajeriarison, E.); Ranomafana N.P., m m (Fisher, B. L.; et al.); Ivohibe, 1500 (Decary, Vohiparara Forest, 1200 (Rajeriarison, E.); R.); Maharira Forest, Ranomafana Natl Pk, 1350 Ranomafana National Park, Talatakely, 915- m m (Rajeriarison, E.);mMaharira Forest, Ranoma- 1000 (Lee, V. F.; Ribrado, K. J.); Ranomafana fana Natl Pk, 1375 (Rajeriarison, E.); Miar- NatlPk. (Rajeriarison,E.);RanomafanaNatlPk., m m anony, Ranomafana Natl Pk, 1050 (Rajeriar- 950-1100 (Bartolozzi, L.; et al.); Ranomafana m ison, E.); Miaranony, Ranomafana Natl Pk, 700 Natl. Pk., Saharoemba ZP, 800 (Rabeson, P.); m (Rajeriarison, E.); Pays de Betsileo, "Sud Ranomafana NP, Talatakely (Griswold, C. E.; Central Madagascar" (Hildebrandt); P.N. Rano- Ubick, D.); Ranomafana, Ambatolahy forest mafana, 0.4 km WSW park entrance, 900 m (Rajeriarison, E.); Ranomafana, Ambatovory m (Kavanaugh, D. H.; Kavanaugh, K. M.); P.N. forest, 1035 (Rajeriarison, E.); Ranomafana, m Ranomafana, 1020 (Harin'Hala, R.); P.N. Miaranony Village (Kingman, A.); Ranomafana, Ranomafana, 1130 m (Harin'Hala, R.); P.N. Vohiparara forest, 1160 m (Rajeriarison, E.); Volume 18, Number 1, 2009 293 m m m Vevembe, 600 (Fisher, B. L.; etal.); Toamasina: (Whitacre, D.); Foret Ivohibe, 650 (Fisher, 10kmSCap Este,5 kmW,20 m (Alpert, G. D.); B. L.; et al.); Fort Dauphin (Alluaud, C); 14 km W Cap Est, Ambato, 100 m (Alpert, G. Manatantely, 100 m (Fisher, B. L.; et al.); P.N. D.); 17kmWAndapa,Res. d'Anjanaharibe-Sud, Andohahela,3.8km113=ESEMahamavo,900m 875 m (Alpert, G. D.); 19 km ESE Maroar\tsetra, (Fisher, B. L.; et al.); P.N. Andohahela, Manam- 250 m (Ward, P. S.); 19 km ESE Maroantsetra, panLhy,5.4 km 113= ESE Mahamavo, 650 m 300 m (Ward, P. S.); 19 km ESE Maroantsetra, (Fisher, B. L.; et al.); PN Andohahela, 275 m 350 m (Ward, P. S.); 1 km SSW Andasibe (Fisher, B. L.; et al.); Res. Andohahela, Mar- (=Perinet), 920 m (Ward, P. S.); 6.3 km S osohy, 600 m (Fisher, B. L.); Vallee d'Ambolo, Ambanizana, Andranobe, 25 m (Fisher, B. L.); Col de Sakalavana (Alluaud, C); province 6.5 km SSW Befingotra, Res. Anjanaharibe-Sud, unknown: "Central Madagascar" (Hildebrandt); 875 m (Fisher, B. L.); 6.9 km NE Ambanizana, "Centre de Madag" (Hildebrandt); "Madag." 1080 m (Fisher, B. L.); 6.9 km NE Ambanizana, (Sikora); "Madagascar Centralis" (Hilde- 650 m (Fisher, B. L.); 8 km ESE Andasibe brandt); "Madagascar" (c.u.); "Madagascar" (=Perinet), 800 m (Ward, P. S.); Ambodiriana, (de Gaulle, J.); "Madagascar" (Grandidier); 125 m (Fisher, B. L.; et al.); Ampasimbe, 450 m "Madagascar" (Sikora); "Madagascar/(S.-E.)" (Betsch, J.-M.); Andasibe (Perinet) (Brooks, R. (Decary, R.). m W.); Andasibe PN, 1025 (Harin'Hala, R.); Worker measurements (n = 13). FiW Anosibe An'ala [as "Nosibe, Village de I'lmer- 1.01-1.44, HL 1.20-1.65, LHT 1.07-1.56, CI ina"] (Sikora); Antongil (Mocquerys); Asanor- eyo, 3 km W Anosibe An'ala (Raharimina, C); 0.77-0.88, FCI 0.15-0.17, REL 0.28-0.36, Bale d'Antongil (c.u.); Betampona, 520 m REL2 0.34-0.43, SI 0.74-0.81, FI 0.29-0.36, (Fisher, B. L.; et al.); Kalalao, 100 m (Fisher, B. PLI 0.50-0.59, PWI 0.40-0.53. L.; et al.); Manakambahiny (Pauly, A.); Mont. Worker diagnosis. With characteristics Akirindro, 7.6km341= NNWAmbinarutelo,600 ofthe T.grandidierigroup (see above);basal m (Fisher, B. L.;etal.);Mont. Anjanaharibe, 18.0 margin of mandible edentate; anterior km21= NNE Ambinanitelo,470 m (Fisher,B. L.; clypeal margin broadly convex and crenu- et al.); Mont. Anjanahmaribe, 19.5 km 27° NNE late, directed forward, not anteroventrally; Ambinanitelo, 1100 (Fisher, B. L.; et al.); head relatively elongate (CI 0.77-0.88); N<o5semyM(aWnagrad,beP.(AlSp.)e;rt,NoGs.yD.)M;aNmnogsaybeM,an1g5a0bem, mlaettearnaoltavliesvp^iraocflemmeosroesoomra,lespsrvoitsribuldeinign (Ward,P.S.);NosyManmgabe,20 (Ward,P.S.); dorsally in the mesopropodeal impression; Nosy Mangabem, 300 (Ward, P. S.); Nosy dorsal face of propodeum broadly convex Mangabe, 3 (Fisher, B. L.; et al.); P.N. Mantadia, 895 m (Ratsirarson, H. }.); Perinet in lateral and posterior viev^s; standing P(NNoyMeasn,aJn.aSr.;a-DNaoyr,d,M.22C5);mP(eFriisnheetr,(BR.osLs.,; eEt.aSl..));; p(i0l.o2s-i0t.y4gemnemralilny slpeanrgsteh;)lodnigstpraiibruetdesdetaase PN Zahamena, 860 m (Fisher, B. L.; et al.); PN follow^s: 1 pair between the frontal carinae, m Zahamena,BesakyRiver, 760 (Fisher,B. L.; et 1 pair on upper half of head, 1 pair on the al.); PN Zahamena, Oribe River, 780 m (Fisher, pronotum, 0-2 pairs on the petiole; 1-2 B. L.; et al.); PN Zahamena, Sahavorondrano pairs on the postpetiole; standing pilosity m River, 765 (Fisher, B. L.; et al.); Saint Marie, scattered on successive abdominal seg- Foret de Kalamlao (Madl); St. Marie (c.u.); ments (gastric segments 1-4); short ap- T10amkpomloN,W21E8nakar(Fai,shReers,.B.AnLd.o;heatheall.a);,T4o2li0arma: pressed to subdecumbent hairs absent or (Fisher, B. L.); 10 km NW Enakara, Res. inconspicuous on most of body; integu- Andohahela, 430 m (Fisher, B. L.); 10 km SSW ment mostly sublucid, with fine coriar- Eminiminy, 750 m (Rajeriarison, E.); 11 km NW ious/puncticulate sculpture; body orange- Enakara,Res. Andohahela, 800m (Fisher, B. L.); brown, appendages lighter; head usually 5 km NNW Isaka-Ivondro, Res. Andohahela, dark brown to brownish-black, but con- 280 m (Ward, P. S.); 5 km WNW Mandiso, Res. colorous with rest of body in some north- m Andohahela, 400 (Ward, P. S.); Col de ern populations (see discussion below); Manangotry, c.30 km N Fort Dauphin, —1000 legs uniformly light orange-brown. 294 Journalof Hymenoptera Research: Festschrift Honoring Roy Snelling Comments. This species is typically Both color forms are here treated as bicolored with a black or darkbrown head conspecific but further studies are needed and the remainder of the body a contrast- to clarify their status. It is possible that ing orange-brown. This allows it to be these color morphs show some degree of distinguished from the other two species, reproductive isolation and/or ecotypic T. inermis and T.—merita, with which it is differentiation. As indicated below, they widely sympatric ^both of these usually appear to be involved in a mimicry have the head more or less concolorous complex with some species of Camponotus. with the mesosoma. Some northern popu- Finally it should be noted that there are lations of T. grandidieri have workers that nine specimens of T. grandidieri in the Forel MHNG are unicolorous orange-brown, however, collection in (Geneva) labeled as and these superficially resemble the other "Typus'' or ''Cotypus'' but most are not two species. They can be recognized true types, because the label data exclude because they lack a tooth on the basal this possibility. These non-types include margin of the mandible (present in T. three males (from Andrangoloaka), one merita) and the metanotal spiracle pro- dealate queen (from Andrangoloaka) and trudes from the mesosoma dorsum in one worker (from "Nosibe, Village de profile (not protruding in T. inermis). The ITmerina"), all with a red 'Typus" label, degree of prominence of the metanotal and an alate queen (Madagascar/Sikora) spiracle varies, however, so itis also useful labeled ''Cotypus''. Only three workers in MHNG to examine head shape, which is more are apparently part of the actual elongate in T. grandidieri (worker CI 0.77- type series of T. grandidieri (there is also a 0.88 versus 0.88-0.97 in T. inermis; see also syntype worker in MCSN). To avoid additional discussion under T. inermis). T. confusion I have designated one of the MHNG grandidieri also overlaps in distribution syntype workers as lectotype. with T. hespera in northern Madagascar. Distribution and biology. Tetraponera Where these two species co-occur T. grand- grandidieri is widespread in eastern Mada- idieri has a bicolored body, while T. hespera gascar, with a distribution that spans the has a unicolored body and contrasting length of the island (Fig. 23). Populations dark bands on the femora. are restricted to rainforest, at elevations At Betampona (17°53'S 49°12'E) Brian ranging from sea level to 1375 m. As a Fisher collected three nest series of T. result of habitat destruction in the low- grandidieri: one (BLF13292) with unico- lands most populations are found at lored workers, a second (BLF13298) with intermediate or higher elevations. Colonies bicolored workers, and a third (BLF13349) usually occupy dead twigs or branches on with both unicolored and bicolored work- the ground, less commonly in the lower ers, in approximately equal proportions. canopy. During field workin Madagascar I The Betampona workers with light and collected thirteennest series ofthis species, dark heads show no obvious differences ofwhich nine were in dead wood and four other than color. The occurrence of both were located in cavities oflive plants: three forms in the same nest is consistent with in stems of tree saplings {Ixora sp., Leea sp. the view that they are conspecific. In and an unidentified plant), and one in a addition, genetic data (>10 kb of se- cavity in a live root of a tree in the genus quence data from several nuclear genes Rhus. There were no scale insects (Coccoi- and one mitochondrial gene) from popu- dea) in any of these live cavity nests, lations sampled throughout the range of however, and there is no indication that the species show the two color forms to T. grandidieri is closely associated with any be phylogenetically comingled (Ward particular plant species. It seems clear that unpubL). it and other members of the T. grandidieri

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