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

2009·13.5 MB·English

by  Philip S Ward

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