Pachycondyla cambouei (Forel)

Rakotonirina, J. C. & B. L. Fisher, 2013, Revision of the Pachycondyla wasmannii - group (Hymenoptera: Formicidae) from the Malagasy region, Zootaxa 3609, pp. 101-141 : 108-119

Pachycondyla cambouei (Forel)

(Figures 6, 8, 10, 14-28, 44-57)

Bothroponera cambouei Forel, 1891 : 133, pl. 4, fig. 7 Lectotype worker, present designation, Madagascar, Imerina, Antananarivo ( Camboué) AntWeb CASENT0101770 (MHNG) [examined]. Paralectotype worker and queens with same data as lectotype but with specimen codes: CASENT0101028, CASENT0101029 (MHNG) and CASENT0101404, CASENT0101402 (MNHN) [examined]. [Combination in Pachycondyla (Bothroponera) : Emery, 1901: 45, 1911: 78; Combination in Bothroponera : Wheeler, 1922: 1007; in Pachycondyla by Bolton, 1995: 303].

Worker diagnosis:

With head in full-face view, antennal scape not surpassing posterior cephalic margin; dorsum of head and mesosoma not finely striate; dorsum of the body with erect slender hairs and pubescence; basal half of antennal scape rounded; dorsal outline of mesosoma not forming a continuous convexity; distinct angle discernable between propodeal dorsum and declivitous surface; petiole nodiform, not anteroposteriorly flattened; anterior half of fourth abdominal tergite (gastral tergite 2) mostly smooth and shiny between large punctures.

Worker measurements (n=116): HL: 1.38-2.29, HW: 1.19-2.01, CI: 85-95, SL: 0.92-1.69, SI: 72-87, PW: 0.95-1.71, WL: 1.64-2.89, NL: 0.65-1.05, NW: 0.79-1.26, NH: 0.95-1.54, DNI: 110-137, LNI: 130-169.

Description:

Worker. In full-face view head widest at or slightly behind level of eyes; sides somewhat convex, but weakly converging in front of eyes; shape of posterior margin variable, ranging from straight to strongly medially excised. Compound eyes either large and moderately protruding, or small with diameter less than half of maximum diameter of scape; with head in full-face view, eyes breaking lateral margins of the head. Antennal scape subcylindrical, with rounded leading edge, and not surpassing posterior margin of head. Anterior clypeal margin truncate and either straight or slightly concave. Mandibles triangular, apical margins armed with seven to nine teeth and denticles. With mesosoma in profile, the dorsal outline not forming a continuous convexity, propodeal dorsum meeting declivity at a distinct angle; mesopleural sulcus usually indistinct and incomplete. Petiole nodiform; in lateral view, anterior face, dorsum, and posterior face meet in a rounded angle. With petiole node in dorsal view, posterior margin straight or medially excised. Hind basitarsus nearly rounded and not flattened dorsoventrally, basal half of inside surface not concave.

Mandibles longitudinally striate and covered with sparse piligerous punctures. Dorsum of head finely reticulate-punctate or reticulate rugulose, superimposed by punctures or setae-bearing foveolae; sides with less defined sculpture, but finely reticulate-punctate or reticulate-rugulose, interspersed with small piligerous pits. Mesosoma and petiole node with variable sculpture, dorsally ranging from coarsely densely punctate to reticulate-rugose or irregularly punctate, interspersed with quite closely spaced smaller punctures; generally fine striations running to bottom of large punctures. For lateral portions, sculpturing varies from finely rugulose to transversely striate with scattered punctures. Propodeal declivity smooth and shiny or with granular surface. Third and fourth abdominal tergites covered with large, shallow punctures, interspaces either smooth and shiny or covered with fine, small punctures; in some specimen s the large punctures are irregularly spaced and much more accentuated. Dorsum of body covered with erect slender or suberect thick hairs, which are reduced in number or absent on the antennal scape and legs. Pubescence on the third and fourth abdominal tergites either dense, sparse, or lacking. Body colors range from ferruginous red to reddish brown through dark brown to black with lighter appendages.

Queen. Measurements (n=14): HL: 1.53-2.18, HW: 1.41-2.09, CI: 89-98, EL: 0.28-0.42, OI: 17-21, SL: 1.02-1.66, SI: 72-82, PW: 1.21-1.87, WL: 2.36-3.45, NL: 0.63-1.02, NW: 0.94-1.34, NH: 0.92-1.35, DNI: 130-153, LNI: 129-146. Winged queens and ergatoids are present in cambouei and its variants. Ergatoid queens and workers look very similar, but the former have ocelli and incomplete thoracic sclerites while the body of alate queens is noticeably larger in general.

Variation:

Pachycondyla cambouei shows a remarkable range of phenotypic diversity. To faciliate discussion of this diversity, the morphological characters of the worker caste are divided into seven forms on the basis of the shape of the occipital corner, the posterior margin of the head, the size of the eyes, the form of the mesosoma, and the abundance of pubescence on the fourth abdominal tergites. However, there is no simple pattern to the degree of variation of these forms. Specifically, a few forms have very restricted geographic boundaries while others show a wide distribution along the humid forests of Madagascar. Each form of P. cambouei is described below:

Form 1 (Figs 10, 15, 18, 22, 24-25, 53). Worker measurements (n=35 workers): HL: 1.75-2.17, HW: 1.59-1.99, CI: 88-95, SL: 1.24-1.57, SI: 75-85, PW: 1.23-1.67, WL: 2.15-2.89, NL: 0.82-1.02, NW: 0.96-1.24, NH: 1.16-1.54, DNI: 110-130, LNI: 130-157.

This form corresponds to the typical P. cambouei , which is distinguished by much larger eyes, medially excised posterior cephalic margin and normal shape of head and mesosoma, medially excised posterior margin of petiole, and smooth or sparsely punctate fourth abdominal tergites. Some characters are variable, such as the smooth or shagreenate propodeal declivity, and slender erect hairs with reduced pubescence or shorter and much more robust erect hairs with more pubescence. A few larger worker specimens (HW: 1.86-1.99 mm) from the Masoala area in northeastern Madagascar are included in this form. These larger workers have less defined sculpture and are covered with elongate and slender erect hairs and less pubescence. At first glance, these differences are suggestive of a distinct species, but other specimens from various localities also show patterns of less enhanced sculpture, slender and long erect hairs, and a larger-sized body than a more typical P. cambouei specimen.

Form 2 (Figs 8, 16, 26-27, 54). Worker measurements (n= 8): HL: 1.83-2.19, HW: 1.66-1.96, CI: 88-91, SL: 1.33-1.51, SI: 77-81, PW: 1.26-1.48, WL: 2.35-2.69, NL: 0.82-0.97, NW: 0.98-1.18, NH: 1.19-1.44, DNI: 114-121, LNI: 138-148.

Known only from workers collected in the isolated rainforest of the RS Ambohijanahary. This variant is characterized by a backward extension into a lobe of each angle of the posterior margin of the head (Fig. 16). However, workers with more typical characters of P. cambouei are also found within a few colonies of this variant, and workers that show intermediate levels of protrusion between this form and form 3 (see below) also occur in FC Didy, FC Andriantantely and PN Mantadia. Workers with intermediate levels of protrusion are included in the latter form.

Form 3 (Figs 6, 17, 28-29, 55). Worker measurements (n=25): HL: 1.92-2.22, HW: 1.72-2.03, CI: 86-93, SL: 1.40-1.58, SI: 77-82, PW: 1.29-1.55, AL: 2.37-3.00, NL: 0.91-0.99, NW: 1.04-1.17, NH: 1.3-1.43, DNI: 112-125, LNI: 140-152.

This variant is distinguished by the fact that the posterolateral margin of the head at each occipital corner protrudes posteriorly into a hornlike tooth. Individual workers with intermediate degrees of expansion of the occipital lobes are known from Ivohibe and PN Mantadia, and this trait varies gradually along its geographical distribution to match the general diagnosis of typical P. cambouei (form 1). Interestingly, worker specimens in some colonies from the PN Ranomafana also present this gradual form of hornlike tooth, which for this form may be indicative of some biological meaning in terms of colony behavior.

Form 4 (Figs 19, 30-31, 54). Worker measurements (n=10): HL: 1.74-1.97, HW: 1.55-1.82, CI: 89-92, SL: 1.25-1.48, SI: 79-86, PW: 1.21-1.38, WL: 2.14-2.55, NL: 0.84-0.99, NW: 0.97-1.14, NH: 1.22-1.34, DNI: 111-126, LNI: 135-151.

Workers of this form are differentiated by the absence of erect hairs and the absence or great reduction in abundance of pubescence on the dorsum of the propodeum. Numerous specimens that show an intermediate abundance of erect hairs and pubescence on the propodeal dorsum also are known across the range of P. cambouei . Specimens of this form could probably be misidentified as P. masoala due to the absence of erect hairs on propodeal dorsum and less defined propodeal sculpture. However, the thickness of the petiolar node, which is about as broad as long when viewed laterally, combined with the gradient of characters, are those of a true P. cambouei .

Form 5 (Figs 20, 32-33, 54). Worker measurements (n=5): HL: 2.10-2.29, HW: 1.82-2.01, CI: 87-88, SL: 1.54-1.69, SI: 83-85, PW: 1.49-1.71, WL: 2.71-2.88, NL: 0.92-1.05, NW: 1.15-1.26, NH: 1.48-1.52, DNI: 118-126, LNI: 141-163.

Two different altitudes in PN Masoala in northeast Madagascar yielded a few workers (4) in which both anterodorsal angles of the pronotum are bidentate or tuberculate where they meet the cervical shield, and the ventral surface of the head near the occipital angles is distinctly compressed; the integument of the entire body is ferruginous red. At first glance this form appears to be a morphologically distinct species, but among the four workers, there is one specimen with a pronotum almost angulate in dorsal view, with less developed teeth, the head apparently not strongly flattened ventrally near the occipital corners. An additional worker collected from the same national park presents a rounded anterodorsal angle of the pronotum when viewed dorsally and rounded occipital angles with the head in profile. The presence of these intermediate characters seems to link this variant to the other forms of P. cambouei . Colonies with all castes of this form should be collected in order to investigate how this character variation relates to the biology of the population and to clarify relationships with the other forms.

Form 6 (Figs 14, 21, 23, 34-35, 56). Worker measurements (n=30 workers): HL: 1.60-2.07, HW: 1.46-1.87, CI: 86-93, SL: 1.11-1.57, SI: 74-87, PW: 1.12-1.47, WL: 2.02-2.74, NL: 0.76-1.02, NW: 0.93-1.18, NH: 1.08-1.41, DNI: 112-135, LNI: 132-169.

This is one of the forms most similar to typical P. cambouei . Workers can be recognized by the combination of the following characters: smaller eyes, straight posterior cephalic margin, slightly concave anterior clypeal margin, flagellar segments as long as broad, shagreened propodeal declivity, and less accentuated sculpture. Abdominal tergite 4 densely covered with appressed hairs when viewed dorsally; petiole node with lateral margins strongly converging anteriorly to form a bluntly rounded tip; posterior margin slightly concave with a weak median excision; body color reddish brown. This form is dominant in the humid, high altitude forests of southern Madagascar.

Form 7 (Figs 36-37, 57). Worker measurements (n=20): HL: 1.38-1.65, HW: 1.19-1.44, CI: 85-92, SL: 0.92-1.13, SI: 72-80, PW: 0.95-1.18, WL: 1.64-2.07, NL: 0.65-0.81, NW: 0.79-1.05, NH: 0.95-1.17, DNI: 119-137, LNI: 139-152.

As in form 6 but workers tend to have smaller body size, broader than long basal flagellar segments, straight anterior clypeal margin, short and robust mesotibiae, and less pubescence on abdominal tergite 4. In dorsal view, petiole node nearly subquadrate, with straight or slightly concave posterior margin and broadly rounded anterior margin; reddish brown to dark brown in color with lighter gaster and appendages. In the north of Madagascar, workers of this form are slightly sculptured and noticeably shiny, with much longer erect hairs, whereas those from the south possess finer, stronger sculpture and shorter hairs. In this variant, morphological diversity occurs and continuously intergrades into that of form 6. Form 7 dominates the rainforests in northern Madagascar but is absent from the PN Montagne d’Ambre.

Discussion:

A better understanding of species boundaries in Malagasy ants is made possible by recent improvements in collecting methods as well as intensive sampling across the entire region. Although morphological differences can distinguish several forms within P. cambouei , enough gradual variation in character states exists across their geographical range to prevent us from hypothesizing separate species for each form. For example, differences in the shape of the posterior margin and the posterior corner of the head, density and size of punctures, and other body sculpture characteristics are not consistent across the region.

Madagascar is characterized by complex topography and habitat heterogeneity. A mountain plateau on the island creates habitats ranging from montane forest at high altitudes and dry forest on its western slopes to lowland rainforest and littoral forests on its eastern side. Continuous environmental gradients exist between each type of habitat. Across these ecological gradients and ecotones, a widespread species like P. cambouei would be expected to show some variation, especially if the suitiable habitats are isolated ( Raxworthy & Nussbaum 1995; Ricklefs 2004; Wiens & Donoghue 2004; Wilmé et al. 2006; Boumans et al. 2007; Smith et al. 2007; Vences et al. 2009). An example of isolated populations is the form 2 located in the relict montane rainforests of RS Ambohijanahary, which is nested in extensive dry habitats in the western slope of Madagascar. The area may have served as a habitat refugia during periods of climatic fluctuations ( Wilmé et al. 2006; Yoder and Heckman 2006; Wollenberg et al. 2008; Pearson and Raxworthy 2009; Vences et al., 2009).

To better understand the morphological diversity of P. cambouei , especially in a historical context, we hope future studies will include molecular sequence data and other morphological data such as male characters which have proven useful in distinguishing cryptic species of ants ( LaPolla et al. 2011). The use of molecular sequence data of P. cambouei will also help evaluate the biogeographical mechanisms that underlie the evolutionary diversification of taxa in Madagascar (as reviewed by Vences et al. 2009).

Distribution and biology:

Pachycondyla cambouei is an endemic, widespread species found from the north throughout the center and the south of Madagascar. It occupies mainly mesic forests at higher and lower altitudes, and also occurs in littoral and transitional forest habitats. This species dominates the representatives of the genus Pachycondyla across its distribution range. Two or more different forms within the species occur sympatrically in several localities; these sympatric groupings include forms 2 and 6; forms 3 and 6; forms 3 and 7; forms 6 and 7; forms 1, 4, and 7; and forms 1, 5, and 7. Although this species is generally terrestrial, a few specimens have been found foraging on low vegetation. Forms 2-7 occur generally on the mountaintops, while forms 1 and 4 also can be found in lowland forests. Most forage through leaf litter and rarely on the forest floor. Nest series were discovered for most forms, collected frequently from rotten logs, under stones, in the ground, in rotten tree stumps, and between root mat layers, but seldom in dead twigs or branches above the ground. Workers of P. cambouei often play dead when their nests are disturbed. Colonies of forms 5 and 6 have not yet been found, which suggests they may be soil dwellers. The smaller eyes, longer erect hairs, and abundant pubescence in form 6 are also suggestive of an underground life style.

Other material examined:

Form 1. MADAGASCAR: Antananarivo: Antananarivo ( Camboué) (MNHN); Tam Perinet; B.M. 1983-201, 27 Apr-3 May 1983 (J.S. Noyes, M.C. Day) (BMNH); Amparihibe (SB); 18; BMNH (E)2003-61, -15.0353, 49.5839 (K.A. Jackson, D. Carpenter) (BMNH); Andrangoloaka (Sikora) (MHNG); Antananarivo; Museum Paris, Grandidier 1893 ( Camboué) (MNHN); Imérina; Museum Paris, Grandidier ( Camboué) (MNHN); Imerina ( Camboué) (MHNG); (W.M. Wheeler) (MCZC); 25 km NNE Ankazobe, -18.1, 47.18333, 1500 m, montane rainforest (P.S. Ward) (MCZC); 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (B.L. Fisher et al.) (CASC); Ankazobe, -18.31617, 47.11583, 1241 m, urban/garden (B.L. Fisher et al.) (CASC); Kaloy, -18.58998, 47.65102, 1423 m, disturbed montane rainforest (B.L. Fisher et al.) (CASC); RNI Sohisika, Sohisika 24.6 km NNE Ankazobe, -18.10322, 47.18692, 1464 m, gallery montane forest (B.L. Fisher et al.) (CASC); RS Ambohitantely, F d’Ambohitantely, 20.9 km 72° NE Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Ambohitantely, F d’Ambohitantely, Jardin Botanique, 24.1 km 59° NE Ankazobe, -18.17139, 47.28182, 1620 m, montane rainforest (Fisher, Griswold et al.) (CASC); Tsinjoarivo, E Ambatolampy (W.L. Brown) (MCZC); Angavokely, -18.93333, 47.75 (B. Pettersson) (PSWC); Vanjamanitra, Antananarivo, 1380 m (J.M. Betsch) (MCZC); Bongolava, Pref. Tsiroanomandidy (A. Peyrieras) (MCZC); Antsiranana: 9.2 km WSW Befingotra, RS Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Ampasindava, F d'Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Ankobahoba, 32.3 km N Ambanja, -13.39166, 48.48249, 41 m, disturbed littoral rainforest (B.L. Fisher et al.) (CASC); Betaolana Forest, along Bekona River, -14.52996, 49.44039, 880 m, rainforest (B.L. Fisher et al.) (CASC); F Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CASC); F d'Anabohazo, 21.6 km 247° WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F de Binara, 9.1 km 233° SW Daraina, -13.26333, 49.60333, 650-800 m, rainforest (B.L. Fisher et al.) (CASC); F de Binara, 9.4 km 235° SW Daraina, -13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher et al.) (CASC); F d'Orangea, 3.6 km 128° SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) (CASC); Marojejy R.N. I. #12, -14.44533, 49.78564, 375 m, rainforest (G.D. Alpert) (MCZC); Marojejy, tributary Manantenina R., - 14.43333, 49.75, 750 m (Quinter & Nguyen) (CASC); Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, - 13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold et al.) (CASC); 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) (CASC); 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) (CASC); 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 et al.) (CASC); PN Montagne d'Ambre, 3.6 km 235° SW Joffreville, -12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Montagne d'Ambre, Antomboka, -12.51269, 49.17807, 970 m, montane rainforest (B.L. Fisher et al.) (CASC); PN Montagne d'Ambre, Crête, -12.58132, 49.13368, 1110 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Manongarivo, 12.8 km 228° SW Antanambao, -13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); Makirovana Forest: -14.1667, 49.95, 715 m; -14.1604, 49.9522, 550 m; -14.1707, 49.9541, 415 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: 29.5 km WNW Tolanaro, Vasiha Mt., -24.93694, 46.75361, 200 m, rainforest (K.C. Emberton et al) (CASC); 45 km S Ambalavao, -22.21667, 47.01667, 720-785 m, rainforest (B.L. Fisher) (CASC); 8.0 km NE Ivohibe, -22.42167, 46.89833, 1200 m, montane rainforest (Sylvain) (CASC); 9.0 km NE Ivohibe, -22.42667, 46.93833, 900 m, rainforest (Sylvain) (CASC); F d'Ambalagoavy Nord, Ikongo, Ambatombe, -21.8275, 47.33889, 625 m (R. Harin'Hala & M.E. Irwin) (CASC); F d'Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CASC); F de Vevembe, 66.6 km 293° Farafangana, -22.791, 47.18183, 600 m, rainforest, transition to montane forest (B.L. Fisher et al.) (CASC); PN Ranomafana, Ambatolahy Forest, montane rainforest (Alpert et al.) (MCZC); PN Andringitra, F Ravaro 12.5 km SW Antanifotsy, -22.21167, 46.845, 1500-1800 m (S. Razafimandimby) (CASC); 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.) (CASC); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Ivohibe: 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (Sylvain) (CASC); RS Ivohibe, 7.5 km ENE Ivohibe, -22.47, 46.96, 900 m, rainforest (Sylvain) (CASC); Ranomafana Nat. Park, Maharira Forest, -21, 47, 1190 m, montane forest (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Miaranony Forest, -21.25, 47.41667, 700 m, montane forest (Emile Rajeriarison) (MCZC); RF d'Agnalazaha, Mahabo, 42.9 km 215° Farafangana, -23.19383, 47.723, 20 m, littoral rainforest (B.L. Fisher et al.) (CASC); RS Manombo 24.5 km 228° Farafangana, -23.01583, 47.719, 30 m, rainforest (B.L. Fisher et al.) (CASC); Mahajanga: 10 km N Mahatsinjo, -17.84028, 47.02472, 1355 m, tropical dry forest (Alpert et al.) (MCZC); RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transition humid forest (B.L. Fisher et al.) (CASC); Toamasina: 33 km south of Moramanga, Rte. d'Anosibe, - 19.25, 48.21667, forest humus and litter (A. Peyrieras) (MCZC); 5.3 km SSE Ambanizana, Andranobe, -15.66667, 49.96667, 425 m, rainforest (B.L. Fisher) (CASC); 6.3 km S Ambanizana, Andranobe, -15.68131, 49.958, 25 m, rainforest (B.L. Fisher) (CASC); 6.9 km NE Ambanizana, Ambohitsitondroina, -15.56667, 50, 825 m, rainforest (B.L. Fisher) (CASC); Ambanizana, PN Masoala, -15.57167, 50.00611, 800-897 m, montane rainforest (D. Andriamalala, D. Silva et al.) (CASC); Ambanizana, PN Masoala, -15.57222, 50.00694, 930-1110 m, montane rainforest (D. Andriamalala, D. Silva et al.) (CASC); Amparihibe (SB), 18 and 31, BMNH (E) 2003-61, [-15.0353, 49.5839] (K.A. Jackson, D. Carpenter) (BMNH); Ambatovy, 12.4 km NE Moramanga, -18.84963, 48.2947, 1010 m, montane rainforest (B.L. Fisher et al.) (CASC); Ambatovy, 12.4 km NE Moramanga, -18.84773, 48.29568, 1000 m, grassland (B.L. Fisher et al.) (CASC); Ambatovy, 12.4 km NE Moramanga, -18.85813, 48.28488, 1040 m, grassland (B.L. Fisher et al.) (CASC); Ambatovy, 12.4 km NE Moramanga, -18.84963, 48.2947, 1010 m, montane rainforest (B.L.Fisher et al.) (CASC); F Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (B.L. Fisher) (CASC) Analalava, 7.0 km 255° Mahavelona, -17.7095, 49.454, 50 m, littoral rainforest (B.L. Fisher et al.) (CASC); Analamay, -18.80623, 48.33707, 1068 m, montane rainforest (Malagasy ant team) (CASC); Beforona, 29 km east of Perinet, -18.96667, 48.58333, 500 m, forest (A. Peyrieras) (MCZC); Bevolota 17.1 km N Andasibe, -18.77071, 48.43164, 995 m, montane rainforest (B.L. Fisher et al.) (CASC); Didyforét (A. Pauly) (CASC); FC Andriantantely, -18.695, 48.81333, 530 m, rainforest (H.J. Ratsirarson) (CASC); FC Sandranantitra, -18.04833, 49.09167, 450 m, rainforest (H.J. Ratsirarson) (CASC); Ile Sainte Marie, F Kalalao, 9.9 km 34° Ambodifotatra, -16.9225, 49.88733, 100 m, rainforest (B.L. Fisher et al.) (CASC); La Mandraka, 1280 m, montane forest (W.L. & D.E. Brown) (MCZC); Mahavelona (Foulpointe), -17.66667, 49.5, forest (A. Pauly) (CASC); Manakambahiny, near Vavatenina Forest, -17.46667, 49.35 (A. Pauly) (CASC); Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, -15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CASC); Morarano-Chrome, 25 km W, forét (A. Pauly) (CASC); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CASC); PN Andasibe-Mantadia, F de Mantadia, 25.7 km 248° Moramanga, -18.81402, 48.43028, 1040 m, rainforest (F.N. Raharimalala, B. Blaimer) (CASC); PN Zahamena, - 17.73359, 48.72625, 950 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Besaky River, -17.75244, 48.85321, 760 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Onibe River, -17.75908, 48.85468, 780 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Sahavorondrano River, -17.75257, 48.85725, 765 m, rainforest (B.L. Fisher et al.) (CASC); PN Zahamena, Tetezambatana Forest, near junction of Nosivola and Manakambahiny Rivers, -17.74298, 48.72936, 860 m, rainforest (B.L. Fisher et al.) (CASC); PN Mananara-Nord, 7.1 km 261° Antanambe, -16.455, 49.7875, 225 m, rainforest (B.L. Fisher et al.) (CASC); Parcelle K7 Tampolo, - 17.28333, 49.41667, 10 m, littoral forest (Malagasy ant team) (CASC); Parcelle K9 Tampolo, -17.175, 49.268, 10 m, littoral forest (Malagasy ant team) (CASC); Perinet (E.S. Ross) (MCZC); Perinet and vic., -18.93333, 48.41667, rainforest (W.L. Brown) (MCZC); Perinet; B.M. 1983-201 (J.S. Noyes, M.C. Day) (BMNH); Res. Ambodiriana, 4.8 km 306° Manompana, along Manompana River, -16.67233, 49.70117, 125 m, rainforest (B.L. Fisher et al.) (CASC); Res. Perinet, rainforest (L.Bartolozzi, S.Tiati & C.Raharimina) (MCZC); RNI Betampona, Camp Rendrirendry 34.1 km 332° Toamasina, -17.924, 49.19967, 390-550 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, -17.88667, 49.2025, 520 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, Betampona 35.1 km NW Toamasina, -17.91801, 49.20074, 500 m, rainforest (B.L. Fisher et al.) (CASC); RNI Betampona, 34.08 km 332° Toamasina, -17.91977, 49.20039, 525 m, rainforest (B.L. Fisher) (CASC); RS Ambatovaky, Sandrangato River, -16.81753, 49.29498, 360 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.77274, 49.26551, 450 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.7755, 49.26427, 430 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.7633, 49.26692, 520 m, rainforest (B.L. Fisher et al.) (CASC); RS Ambatovaky, Sandrangato River, -16.76912, 49.26704, 475 m, rainforest (B.L. Fisher et al.) (CASC); SF Tampolo, 10 km NNE Fenoarivo Atn., -17.2825, 49.43, 10 m, littoral rainforest (B.L. Fisher) (CASC); Sahafina forest 11.4 km W Brickaville, -18.81445, 48.96205, 140 m, rainforest (B.L. Fisher et al.) (CASC); SF Analamazaotra, Analamazaotra 1.3 km S Andasibe, -18.38466, 48.41271, 980 m, montane rainforest (B.L. Fisher et al.) (CASC); Torotorofotsy, -18.87082, 48.34737, 1070 m, montane rainforest, marsh edge (Malagasy ant team) (CASC); Torotorofotsy, -18.87467, 48.3725, 960 m, rainforest (Woodhead & Vences) (CASC); 1 km W Andampibe, Cap Masoala, -15.69361, 50.18139, 125 m, lowland forest (G.D. Alpert) (MCZC); Rés. Ambodiriana 45 km NW Toamasina (P. Rabeson) (MCZC); vic Andasibe (=Perinet), 950 m, forest (W.L. & D.E. Brown) (MCZC); Toliara: 29.5 km WNW Tolanaro, Vasiha Mt., -24.92306, 46.74083, 700 m, rainforest (K.C. Emberton et al.) (CASC); 10 km NW Enakara, Rés Andohahela, -24.56667, 46.81667, 420 m, rainforest (B.L. Fisher) (CASC); 11 km NW Enakara, Rés. Andohahela, -24.56667, 46.83333, 800 m, rainforest (B.L. Fisher) (CASC); 13 km NW Enakara, Rés. Andohahela, -24.55, 46.8, 1250 m, montane rainforest (B.L. Fisher) (CASC); 13 km NW Enakara, Rés Andohahela, -24.55, 46.8, 1280 m, montane rainforest (B.L. Fisher) (CASC); 2.7 km WNW 302° Ste. Luce, - 24.77167, 47.17167, 20 m, littoral rainforest (J.-Baptiste) (CASC); 6 km SSW Eminiminy, Res. Andohahela, - 24.73333, 46.8, 330 m, rainforest (E. Rajeriarison) (MCZC); Andohahela, -24.77639, 46.70528, 320 m, tropical dry forest (P. Rabeson) (MCZC); Andohahela, Parcel #1 versante E., 300 m, rainforest (L. Bartolozzi, S. Tiati & C. Raharimina) (MCZC); Res. Andohahela, 6 km ESE Imonty, -24.84444, 46.75361, 1000 m, rainforest (P. Rabeson) (MCZC); Res.Andohahela, 9 km SSW Eminiminy, -24.75, 46.78333, 500 m, rainforest (E. Rajeriarison) (MCZC);

FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); FC Analavelona, 33.2 km 344° NNW Mahaboboka, -22.64333, 44.17167, 1300 m, montane rainforest (Fisher, Griswold et al.) (CASC); F Ivohibe 55.0 km N Tolagnaro, -24.569, 47.204, 200 m, rainforest (B.L. Fisher et al.) (CASC); Manatantely, 8.9 km NW Tolagnaro, -24.9815, 46.92567, 100 m, rainforest (B.L. Fisher et al.) (CASC); PN Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, -24.7585, 46.85367, 275 m, rainforest (B.L. Fisher et al.) (CASC); 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) (CASC); PN d'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) (CASC); Res. Andohahela, 5 km SSW Eminiminy, - 24.73333, 46.8, 200 m (E. Rajeriarison) (MCZC); RS Ambohijanahary, F d'Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala, -18.26, 45.41833, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Ambohijanahary, F d'Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, -18.26667, 45.40667, 1050 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Kalambatritra, Ambinanitelo, -23.4502, 46.45658, 1325 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.4635, 46.4631, 1270 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.463, 46.47057, 1269 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.4635, 46.4631, 1270 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.463, 46.47057, 1269 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Befarara, -23.4178, 46.4478, 1390 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Betanana, -23.4144, 46.459, 1360 m, montane rainforest (B.L. Fisher et al.) (CASC).

Form 2. Toliara: RS Ambohijanahary, F d'Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, - 18.26667, 45.40667, 1050 m, montane rainforest (Fisher, Griswold et al.) (CASC). Form 3. Antsiranana: PN Montagne d'Ambre [1st campsite], -12.51444, 49.18139, 960 m, rainforest (M.E. Irwin, E.I. Schlinger, R. Harin'Hala) (CASC); PN Montagne d'Ambre [Petit Lac road], -12.52028, 49.17917, 1125 m, rainforest (R. Harin'Hala) (CASC); Fianarantsoa: 14 km W of park headquarters, Ranomafana National Park, - 21.22707, 47.36874, 1100 m, cloud forest (M.E. Irwin & E.I. Schlinger) (CASC); 2 km W Andrambovato, along river Tatamaly, -21.51167, 47.41, 1075 m, montane rainforest (B.L. Fisher et al.) (CASC); 3 km W Ranomafana, nr. Ifandiana, -21.25, 47.41667, 950 m, forest (P.S. Ward) (PSWC); 3 km WNW Ranomafana, nr. Ifanadiana, - 21.25, 47.43333, 840 m, roadside (P.S. Ward) (PSWC); 45 km S. Ambalavao, -22.21667, 47.01667, 785 m, rainforest (B.L. Fisher) (CASC); 7.6 km 122 º Kianjavato, FC Vatovavy, -21.4, 47.94, 175 m, rainforest (B.L. Fisher et al.) (CASC); 7 km W Ranomafana, 1100 m (C. Kremen, R. Van Epps & L. Rasabo) (MCZC); 7 km W Ranomafana, -21.26667, 47.41667, 1000 m, rainforest (M. Stebbins, W.E. Steiner et al.) (MCZC); 9.0 km NE Ivohibe, -22.42667, 46.93833, 900 m, rainforest (Sylvain) (CASC); Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov., -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin'Hala) (CASC); Isalo Nat. Park, Ranohira, Canyon de Sinze, -22.48333, 45.55, 800 m, gallery forest (E. Rajeriarison) (MCZC); Nat. Park Ranomafana, Ambatolahy Forest (E. Rajeriarison) (MCZC); PN Ranomafana (L. Bartolozzi, S. Tiati, & C. Raharimina) (MCZC); PN Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, -23.83517, 46.96367, 940 m, rainforest (B.L. Fisher et al.) (CASC); 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.) (CASC); PN Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, -21.23667, 47.39667, 1150 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Ranomafana Talatakely, -21.24833, 47.425, 900 m, between main bridge and park entrance (J.S. Schweikert) (CASC); PN Ranomafana: Talatakely, -21.24833, 47.42667, in guava forest (C.E. Griswold, D.H. Kavanaugh, N.D. Penny, M.J. Raherilalao, J.S. Ranorianarisoa, J.S. Schweikert) (CASC); P.N. Ranomafana: Talatakely, -21.24833, 47.42667, in bamboo forest (C.E. Griswold, D.H. Kavanaugh, N.D. Penny, M.J. Raherilalao, J.S. Ranorianarisoa, J.S. Schweikert) (CASC); RS Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (Sylvain) (CASC); RS Ivohibe, 6.5 km ESE Ivohibe, - 22.49667, 46.955, 1575 m, montane rainforest (Sylvain) (CASC); RS Ivohibe, 7.5 km ENE Ivohibe, -22.47, 46.96, 900 m, rainforest (Sylvain) (CASC); radio tower, Ranomafana National Park, Fianarantsoa Prov., -21.25083, 47.40717, 1130 m, forest edge, mixed tropical forest, open area (M. Irwin, R. Harin'Hala) (CASC); Ranomafana, - 21.25, 47.36667 (A. Pauly) (CASC); Ranomafana (A. Kingman) (MCZC); Ranomafana Nat. Park, 700 m, rainforest (W.E. Steiner) (MCZC); Ranomafana Nat. Park, 7 km W, 900 m, montane rainforest (W.E. Steiner) (MCZC); Ranomafana Nat. Park, Fivondronana Ifanadiana Talatakely, 1100 m (J.E. Cadle) (MCZC); Ranomafana Nat. Park, near Andranoroa, low tree forest (I. Constable) (MCZC); Ranomafana Nat. Park, near research cabin, - 21.25, 47.41667 (V.P. Lee, K.J. Rivardo, leg.) (CASC); Ranomafana Nat. Park, SE research cabin, montane rainforest (P. Rabeson) (MCZC); Ranomafana Nat. Park, Talatakely Forest, montane rainforest (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Valoloaka Forest, 1200 m, montane forest (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Vohiparara, 1160 m, montane rainforest (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Vohiparara Sahamalaotra, -21.24444, 47.39694, rainforest (P. Rabeson) (MCZC); Ranomafana Nat. Park, Mahirira Forest, -21, 47, 1350 m (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Miaranony Forest, -21.25, 47.41667, 800 m, montane forest (E. Rajeriarison) (MCZC); Ranomafana, Miaranony village (A. Kingman) (MCZC); research cabin at Talatakely, Ranomafana National Park, -21.25041, 47.41945, 900 m, mixed tropical forest (M.E. Irwin & E.I. Schlinger) (CASC); Vohiparara broken bridge, Fianarantsoa Prov., -21.22617, 47.36983, 1110 m, high altitude rainforest (R. Harin'Hala) (CASC); Vohiparara, 13 km W Ranomafana, foresta secondaria (L. Bartolozzi, S. Tiati, & C. Raharimina) (MCZC); Toamasina: 19 km ESE Maroantsetra, -15.48333, 49.9, 350 m, rainforest (P.S. Ward) (PSWC); 1 km SSW Andasibe, -18.93333, 48.41667, 420 m, forest (P.S. Ward) (MCZC); 1 km SSW Andasibe, -18.93333, 48.41667, 420 m, forest (P.S. Ward) (PSWC); 1 km SSW Andasibe (=Perinet) -18.93333, 48.41667, 920 m, rainforest (P. S. Ward) (MCZC); 6 km ESE Andasibe (=Perinet), -18.95, 48.46667, 900 m, rainforest (P.S. Ward) (MCZC); 6 km ESE Andasibe(- Périnet), -18.95, 48.46666667, 900 m, rainforest (P.S. Ward) (PSWC); Andranoaria, 15 km S Anosibe An 'alam. (C. Raharimina) (MCZC); FC Andriantantely, -18.695, 48.81333, 530 m, rainforest (H.J. Ratsirarson) (CASC); FC Didy, -18.19833, 48.57833, 960 m, rainforest (H.J. Ratsirarson) (CASC); FC Sandranantitra, -18.04833, 49.09167, 450 m, rainforest (H.J. Ratsirarson) (CASC); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CASC); Perinet and vic., -18.93333, 48.41667, rainforest (W.L. Brown) (MCZC); Perinet and vic., -18.93333, 48.41667, rainforest (W.L. Brown) (MCZC); Res. Perinet, Andasibe, 870 m (Alpert et al.) (MCZC); vic Andasibe (=Perinet), 950 m (W.L. Brown) (MCZC); Toliara: F Ivohibe 55.6 km N Tolagnaro, -24.56167, 47.20017, 650 m, rainforest (B.L. Fisher et al.) (CASC); PN Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, -24.7585, 46.85367, 275 m, rainforest (B.L. Fisher et al.) (CASC).

Form 4. Antsiranana: 6.5 km SSW Befingotra, RS Anjanaharibe-Sud, -14.75, 49.5, 875 m, rainforest (B.L. Fisher) (CASC); 9.2 km WSW Befingotra, RS Anjanaharibe-Sud, -14.75, 49.46667, 1280 m, montane rainforest (B.L. Fisher) (CASC); 9.2 km WSW Befingotra, RS Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Makirovana Forest: -14.1651, 49.9477, 900 m, montane rainforest; -14.1604, 49.9522, 550 m, rainforest (B.L. Fisher et al.) (CASC); Toamasina: Montagne d'Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, -15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, -15.17833, 49.635, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CASC).

Form 5. Toamasina: 6.9 km NE Ambanizana, Ambohitsitondroina, -15.56667, 50, 825 m, rainforest (B.L. Fisher) (CASC); 5.3 km SSE Ambanizana, Andranobe, -15.66667, 49.96667, 425 m, rainforest (B.L. Fisher) (CASC).

Form 6. Antananarivo: 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, -18.47333, 47.96, 1300 m, montane rainforest (Fisher, Griswold et al.) (CASC); 2 km W Ivato (G.D. Alpert) (MCZC); Ambohidratrimo, - 18.821, 47.44183, 1362 m, urban garden (B.L. Fisher et al.) (CASC); F de galerie, Andranorovitra, 24.0 km NNE Ankazobe, -18.11243, 47.19757, 1491 m, disturbed gallery montane forest (B.L. Fisher et al.) (CASC); F de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, -18.12167, 47.20627, 1520 m, disturbed gallery montane forest (B.L. Fisher et al.) CASC); Ilafy, -18.85415, 47.56575, 1385 m, urban garden (B.L. Fisher et al.) (CASC); RNI Sohisika, Sohisika 24.6 km NNE Ankazobe, -18.10322, 47.18692, 1464 m, gallery montane forest (B.L. Fisher et al.) (CASC); RS Ambohitantely, F d’Ambohitantely, 20.9 km 72° NE Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Ambohitantely, F d’Ambohitantely, Jardin Botanique, 24.1 km 59° NE d Ankazobe, -18.17139, 47.28182, 1620 m, montane rainforest (Fisher, Griswold et al.) (CASC); Antsiranana: Bemanevika, 10 km N de Sambava souspref. de Bealanana, -14.13333, 50.11667 (A. Peyrieras) (MCZC); 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 et al.) (CASC); PN montagne d'Ambre, 3.6 km 235° SW Joffreville, - 12.53444, 49.1795, 925 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Fianarantsoa: 2 km W Andrambovato, along River Tatamaly, -21.51167, 47.41, 1075 m, montane rainforest (B.L. Fisher et al.) (CASC); 43 km S Ambalavao, Rés. Andringitra, -22.23333, 47, 825 m, rainforest (B.L. Fisher) (CASC); 7.6 km 122 º

Kianjavato, FC Vatovavy, -21.4, 47.94, 175 m, rainforest (B.L. Fisher et al.) (CASC); 7 km W Ranomafana, 1000 m, rainforest (W.E. Steiner) (MCZC); 8 km E Kianjavato, Vatovavy Forest, -21.39306, 47.94194, 170 m, lowland forest (Alpert et al.) (MCZC); 9.0 km NE Ivohibe, -22.42667, 46.93833, 900 m, rainforest (Sylvain) (CASC); Belle Vue trail, Ranomafana National Park, -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin'Hala) (CASC); F d'Ambalagoavy Nord, Ikongo, Ambatombe, -21.8275, 47.33889, 625 m (R. Harin'Hala & M.E. Irwin) (CASC); F d'Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CASC); F de Vevembe, 66.6 km 293° Farafangana, -22.791, 47.18183, 600 m, rainforest, transition to montane forest (B.L. Fisher et al.) (CASC); Isalo Nat. Park, Ranohira, Canyon de Sinze, -22.48333, 45.55, 800 m, gallery forest (E. Rajeriarison) (MCZC); PN Andringitra, F Ravaro 12.5 km SW Antanifotsy, -22.21167, 46.845, 1500-1800 m (S. Razafimandimby) (CASC); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); Parc naturel communautaire, 28.1 km SW Ambositra, -20.78493, 47.17328, 1727 m, disturbed montane rainforest along roadside (B.L. Fisher et al.) (CASC); RS Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (Sylvain) (CASC); RS Ivohibe, 7.5 km ENE Ivohibe, -22.47, 46.96, 900 m, rainforest (Sylvain) (CASC); Ranomafana Nat. Park, Talatakely Forest, montane rainforest (E. Rajeriarison) (MCZC); Ranomafana Nat. Park, Miaranony Forest, -21.25, 47.41667, 800 m, montane forest (E. Rajeriarison) (MCZC); RS Manombo 24.5 km 228° Farafangana, -23.01583, 47.719, 30 m, rainforest (B.L. Fisher et al.) (CASC); Mahajanga: RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transition humid forest (B.L. Fisher et al.) (CASC); Toamasina: 16 km S Moramanga, -19.08333, 48.23333, 950 m, rainforest (P.S. Ward) (PSWC); 6.9 km NE Ambanizana, Ambohitsitondroina, -15.56667, 50.0095, 825 m, rainforest (B.L. Fisher) (CASC); Ambanizana, PN Masoala, - 15.57167, 50.00611, 900-950 m, montane rainforest (D. Andriamalala, D. Silva et al.) (CASC); FC Andriantantely, -18.695, 48.81333, 530 m, rainforest (H.J. Ratsirarson) (CASC); FC Didy, -18.19833, 48.57833, 960 m, rainforest (H.J. Ratsirarson) (CASC); F Ambatovy, 14.3 km 57° Moramanga, -18.85083, 48.32, 1075 m, montane rainforest (Malagasy ant team) (CASC); F d'Analava Mandrisy, 5.9 km 195 Antanambe, -16.48567, 49.847, 10 m, littoral rainforest (B.L. Fisher et al.) (CASC); Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, -15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CASC); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CASC); PN Marojejy, - 14.43817, 49.774, 488 m, rainforest (Rin'Ha, Irwin) (CASC); RNI Betampona, Camp Rendrirendry 34.1 km 332° Toamasina, -17.924, 49.19967, 390 m, rainforest (B.L. Fisher et al.) (CASC); SF Analamazaotra, Analamazaotra 1.3 km S Andasibe, -18.38466, 48.41271, 980 m, montane rainforest (B.L. Fisher et al.) (CASC); Torotorofotsy, - 18.87082, 48.34737, 1070 m, montane rainforest, marsh edge (Malagasy ant team) (CASC); Toliara: 11 km NW Enakara, Rés. Andohahela, -24.56667, 46.83333, 800 m, rainforest (B.L. Fisher) (CASC); 13 km NW Enakara, Res. Andohahela, -24.55, 46.8, 1250 m, montane rainforest (B.L. Fisher) (CASC); FC Analavelona, 29.2 km 343° NNW Mahaboboka, -22.675, 44.19, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); FC Analavelona, 29.4 km 343° NNW Mahaboboka, -22.675, 44.18667, 1050 m, montane rainforest (Fisher, Griswold et al.) (CASC); F de Petriky, 12.5 km W 272° Tolagnaro, -25.06167, 46.87, 10 m, littoral rainforest (B.L. Fisher) (CASC); Mandena, 8.4 km NNE 30° Tolagnaro, -24.95167, 47.00167, 20 m, littoral rainforest (B.L. Fisher) (CASC); PN Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, -24.7585, 46.85367, 275 m, rainforest (B.L. Fisher et al.) (CASC); 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) (CASC); RS Ambohijanahary, F d'Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, -18.26667, 45.40667, 1050 m, montane rainforest (Fisher, Griswold et al.) (CASC); RS Kalambatritra, Ambinanitelo, -23.4502, 46.45658, 1325 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.4635, 46.4631, 1270 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Ampanihy, -23.463, 46.47057, 1269 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Kalambatritra, Befarara, -23.4178, 46.4478, 1390 m, montane rainforest (B.L. Fisher et al.) (CASC).

Form 7. Antsiranana: 6.5 km SSW Befingotra, RS. Anjanaharibe-Sud, -14.75, 49.5, 875 m, rainforest (B.L. Fisher) (CASC); 9.2 km WSW Befingotra, RS. Anjanaharibe-Sud, -14.75, 49.46667, 1200 m, montane rainforest (B.L. Fisher) (CASC); Ambondrobe, 41.1 km 175° Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest (B.L. Fisher) (CASC); Bemanevika, 10 km N de Sambava souspref. de Bealanana, -14.13333, 50.11667 (A. Peyrieras) (MCZC); F de Binara, 9.4 km 235° SW Daraina, -13.26333, 49.6, 1100 m, montane rainforest (B.L. Fisher et al.) (CASC); 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 et al.) (CASC); RS Manongarivo, 12.8 km 228° SW Antanambao, - 13.97667, 48.42333, 780 m, rainforest (B.L. Fisher) (CASC); RS Manongarivo, 14.5 km 220° SW Antanambao, - 13.99833, 48.42833, 1175 m, montane rainforest (B.L. Fisher) (CASC); Ambondrobe, 41.1 km 175° Vohemar, - 13.71533, 50.10167, 10 m, littoral rainforest (B.L. Fisher) (CASC); Makirovana Forest, -14.1707, 49.9541, 415 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: 9.0 km NE Ivohibe, -22.42667, 46.93833, 900 m, rainforest (Sylvain) (CASC); RS Ivohibe 8.0 km E Ivohibe, -22.48333, 46.96833, 1200 m, montane rainforest (Sylvain) (CASC); PN Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, -21.29, 47.43333, 1100 m, montane rainforest (Fisher, Griswold et al.) (CASC); Belle Vue trail, Ranomafana National Park, -21.2665, 47.42017, 1020 m, mixed tropical forest (R. Harin'Hala) (CASC); Mahajanga: RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transitional humid forest (B.L. Fisher et al.) (CASC); Toamasina: 6.9 km NE Ambanizana, Ambohitsitondroina, -15.56667, 50.000, 825 m, rainforest (B.L.Fisher) (CASC); Ambanizana, PN Masoala, -15.57167, 50.00611, 900-950 m, montane rainforest (D. Andriamalala, D. Silva et al.) (CASC); FC Andriantantely, -18.695, 48.81333, 530 m, rainforest (H.J. Ratsirarson) (CASC); FC Didy, -18.19833, 48.57833, 960 m, rainforest (H.J. Ratsirarson) (CASC); FC Sandranantitra, -18.04833, 49.09167, 450 m, rainforest (H.J. Ratsirarson) (CASC); Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo, -15.28833, 49.54833, 600 m, rainforest (Fisher, Griswold et al.) (CASC); Montagne d'Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, - 15.18833, 49.615, 470 m, rainforest (Fisher, Griswold et al.) (CASC); PN Mantadia, -18.79167, 48.42667, 895 m, rainforest (H.J. Ratsirarson) (CASC); PN Zahamena, Onibe River, -17.75908, 48.85468, 780 m, rainforest (B.L. Fisher et al.) (CASC); Toliara: 11 km NW Enakara, Rés. Andohahela, -24.56667, 46.83333, 800 m, rainforest (B.L. Fisher) (CASC); 13 km NW Enakara, Res. Andohahela, -24.55, 46.8, 1250 m, montane rainforest (B.L. Fisher) (CASC); 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) (CASC); 0.7 km WNW 302° Ste. Luce, -24.77167, 47.17167, 20 m, littoral rainforest (J.-Baptiste) (CASC); F de Petriky, 12.5 km W 272° Tolagnaro, -25.06167, 46.87, 10 m, littoral rainforest (B.L. Fisher) (CASC); Mandena, 8.4 km NNE 30° Tolagnaro, -24.95167, 47.00167, 20 m, littoral rainforest (B.L. Fisher) (CASC).