Carebara grandidieri (Forel, 1891)

Azorsa, Frank & Fisher, Brian L., 2018, Taxonomy of the ant genus Carebara Westwood (Formicidae, Myrmicinae) in the Malagasy Region, ZooKeys 767, pp. 1-149 : 1

Carebara grandidieri (Forel, 1891)

Oligomyrmex grandidieri Forel, 1891: 201. Holotype queen (1 queen, CASENT0101999), (MHNG) [examined]: Madagascar, Antananarivo ( Camboué). Combination in Carebara : Fernández, 2004: 235.

Oligomyrmex voeltzkowi Forel, 1907: 77. Holotype queen (1 queen, CASENT0906665, GBIF-D/FoCol1927), (ZMHB): Madagascar, Toamasina, Tamatave (Voeltzkow). [examined] syn. n. Combination in Carebara : Fernández, 2004: 235.

Diagnosis.

Antennae nine-segmented. Major: Head longer than wide, nearly rectangular, margins straight and parallel; petiole with relatively long peduncle, ventral face flat, petiolar node rounded. Minor: Head longer than wide, nearly subrectangular; petiole with relatively long peduncle, petiolar node rounded.

Description of major workers.

Measurements (n=22): HL 0.56-0.84; HW 0.44-0.61; SL 0.25-0.32; ML 0.12-0.21; EL 0.01-0.03; EM 0.14-0.19; HD 0.30-0.45; WL 0.51-0.74; PSL 0.05-0.09; PW 0.26-0.38; MFL 0.29-0.42; MFW 0.06-0.10; MTL 0.26-0.37; PTL 0.18-0.28; PNL 0.09-0.14; PTH 0.12-0.20; PTW 0.12-0.19; PPL 0.11-0.16; PPNL 0.09-0.14; PPH 0.10-0.17; PPW 0.13-0.21; GL 0.44-0.85; GW 0.35-0.65; CI 71-79; MI 20-27; SI 38-46; MLI 63-74; PPLI 50-67; PPI 100-123; PSI 10-15.

Head longer than wide (CI 71-79), in full-face view nearly subrectangular, about 1.3 times longer than wide. Posterior margin of head medially concave, posterolateral corners rounded, lateral margins straight. Mandibles with six teeth. Anterior margin of clypeus concave and laterally convex. Frons without median ocellus. Antennae with nine segments. Scapes short (HL 0.56-0.84, SL 0.25-0.32, SI 38-46). Eyes present, consisting of one to three ommatidia (EL 0.01-0.03). Supraclypeal area acutely triangular, extending forward beyond the antennal insertions.

In profile view, promesonotum convex, mesonotum shallowly present and lower than dorsum of promesonotum, metanotal groove present. Propodeum lower than promesonotum, about 1.2 times higher than long, dorsal face of propodeum nearly flat and declining posteriorly, posterodorsal corner angulate to dentate, declivity of propodeum slightly concave with thin lateral laminae in direction of propodeal lobe. Propodeal lobes short and convex. Propodeal spiracle rounded and situated above mid-height of sclerite, and beyond mid-length of sclerite by about half the diameter of the spiracle; distance from propodeal spiracle to posterodorsal corner of propodeum about 2.5 times the diameter of the spiracle (PSL 0.05-0.09), and distance to declivity almost twice the diameter of the spiracle. In dorsal view, promesonotum about 1.2 times longer than wide, anterior margin of promesonotum rounded, sides convex and narrowed posteriorly; sides of propodeum weakly convex or straight.

Petiole with relatively long peduncle, ventral face flat. Combined outline of dorsal surface of peduncle and anterior face of node deeply concave in the middle, posterior face of node slightly convex, anterodorsal and posterodorsal corner convex, dorsum rounded. Subpetiolar process produced as a small denticle, smaller than the diameter of the propodeal spiracle. Postpetiolar node rounded and lower than petiolar node. In dorsal view, petiolar node as broad as postpetiolar node (PTW 0.12-0.19, PPW 0.13-0.21) and petiolar node wider than long (PNL 0.09-0.14, PTW 0.12-0.19), anterior and posterior margins of petiole nearly straight, anterior margin of postpetiole concave and convex posterior margin, sides rounded in petiole and convex in postpetiole, narrowed posteriorly.

Dorsal surface of mandibles, upper median portion of clypeus and supraclypeal area smooth and shiny, with scattered piligerous punctae on head and mandibles. Head with longitudinal rugae along the gena, and frontal lobes. Lower median portion of clypeus with transverse rugae. In profile, posterolateral portion of cephalic dorsum smooth and shiny. Mesosoma smooth and shiny, except for propleuron and mesopleuron (areolate), and metapleuron (longitudinally rugulose). Ventral face of petiole and postpetiole longitudinally areolate. In dorsal view mesosoma, petiole, postpetiole and gaster smooth and shiny, except for declivity of propodeum, anterior and posterior face of petiole finely areolate.

Lateral margins and posterior margin of head with suberect to subdecumbent hairs. Scapes with subdecumbent to decumbent hairs. Outer margin of mandibles with short decumbent hairs. Mesosoma with short and long suberect hairs. Petiole and postpetiole with short subdecumbent hairs and long suberect hairs. Tibia with subdecumbent to decumbent hairs. Gaster with abundant suberect to decumbent hairs. Color yellowish ferruginous.

Description of minor workers.

Measurements (n=18): HL 0.39-0.49; HW 0.31-0.40; SL 0.23-0.30; ML 0.08-0.13; EL 0.01-0.02; EM 0.11-0.13; HD.022-0.28; WL 0.40-0.52; PSL 0.03-0.05; PW 0.18-0.24; MFL 0.24-0.31; MFW 0.06-0.08; MTL 0.19-0.27; PTL 0.16-0.19; PNL 0.09-0.11; PTH 0.10-0.13; PTW 0.09-0.11; PPL 0.08-0.11; PPNL 0.07-0.10; PPH 0.07-0.10; PPW 0.10-0.12; GL 0.31-0.55; GW 0.22-0.32; CI 79-86; MI 19-28; SI 57-65; MLI 73-82; PPLI 47-63; PPI 100-120; PSI 9-13.

Head longer than wide (CI 79-86), in full-face view nearly subrectangular, about 1.2 times longer than wide. Posterior margin of head weakly concave, nearly straight, posterolateral corners rounded, lateral margins slightly convex, nearly straight. Mandibles with five teeth. Anterior margin of clypeus concave and laterally angulate. Antennae with nine segments. Scape fails to reach the posterior margin of head (HL 0.39-0.49, SL 0.23-0.30, SI 57-65). Eyes present, consisting of one ommatidium (EL 0.01-0.02). Supraclypeal area triangular but poorly defined.

In profile view, promesonotum weakly convex, nearly flat, metanotal groove present. Propodeum about 1.3 times higher than long, dorsal face of propodeum convex, posterodorsal corners angulate, or with a small triangular tooth, anterodorsal corner convex, declivity concave with thin lateral laminae in direction of propodeal lobe. Propodeal lobes short and triangular. Propodeal spiracle rounded and situated above mid-height of sclerite by about two times the diameter of the spiracle, and beyond mid-length of sclerite by about 1.5 times the diameter of the spiracle, distance from propodeal spiracle to posterodorsal corner of propodeum almost three times the diameter of the spiracle (PSL 0.03-0.05), and distance to declivity less than twice the diameter of the spiracle. In dorsal view, promesonotum about 1.25 times longer than wide, anterior margin rounded, sides convex; sides of propodeum straight.

Petiole with a relatively long peduncle, petiole longer than high (PTL 0.16-0.19, PTH 0.10-0.13), ventral face flat. Combined outline of dorsal surface of peduncle and anterior face of node deeply concave in the middle, posterior face of node slightly convex, anterodorsal and posterodorsal corner convex, dorsum rounded. Subpetiolar process produced as a small denticle, smaller than the diameter of the propodeal spiracle. Postpetiolar node strongly convex and lower than petiolar node. In dorsal view, petiolar node almost as broad as postpetiolar node (PTW 0.09-0.11, PPW 0.10-0.12), and petiolar node as broad as long (PNL 0.09-0.11, PTW 0.09-0.11), anterior and posterior margin of petiole convex, anterior margin of postpetiole concave and convex posteriorly, sides of petiole and postpetiole strongly convex.

Dorsal surface of mandibles, clypeus, supraclypeal area, and head smooth and shiny, with scattered piligerous punctae on head and mandibles. Head with transverse rugae near the posterior margin of head, gena and frontal lobes with longitudinal rugae. Mesosoma smooth and shiny, except for propleuron and mesopleuron (areolate), and metapleuron (finely areolate-rugose). Lateral margins of petiole finely areolate-rugose, except for petiolar node and ventral face of postpetiole which are areolate. In dorsal view mesosoma, petiole, postpetiole and gaster smooth and shiny, except for declivity of propodeum, anterior and posterior face of petiole which are finely areolate.

Lateral margins of head and scape with subdecumbent to decumbent hairs. Posterior margin of head with short suberect hairs. Outer margin of mandibles with decumbent hairs. Mesosoma with short and long suberect hairs. Petiole and postpetiole with short decumbent hairs and long subdecumbent hairs. Tibia with decumbent hairs. Gaster with longer suberect to decumbent hairs. Color yellowish ferruginous.

Distribution and biology.

Carebara grandidieri is a widespread and common species in the Malagasy region, and has been recorded on Madagascar, Comoros and Mayotte (Figure 68). This species was collected in the following environments: along the roadside, coastal scrub, dry forest, gallery forest, grassland, littoral forest, montane rainforest, open secondary vegetation, rainforest, spiny forest/thicket, tropical dry forest, Uapaca woodland, and urban/garden. C. grandidieri was sampled at elevations ranging from 10 m to 1550 m. Individuals and colonies were collected in rotten logs, rotting tree stumps, soil, termite mounds, leaf litter, under moss, litter on rocks, under rotten logs, under stones, and in leaf litter.

Comments.

Carebara grandidieri is widely distributed throughout Madagascar, Comoros and Mayotte, and co-occurs with almost all Carebara species recorded in the Malagasy region. The morphology of this species is highly variable (Figure 39, 40). There are different populations that initially appear to be distinct species. However, we could not find consistent characters to split C. grandidieri into additional species. The characters that vary across populations include: a) pilosity that varies from suberect to decumbent hairs, and is more noticeable on the gaster; b) posterior corner of propodeum, which varies from a pair of small triangular and angulate upwardly directed teeth, to nearly convex (in specimens from the same colony it is possible to see the gradual change from nearly convex to slightly dentate); c) petiole, where the node is thicker in profile and dorsal view, in some populations; and d) the diameter of the propodeal spiracle, which is smaller in populations from the north of Madagascar.

Upon examining all Carebara workers in the Malagasy region, we found only two species with nine antennal segments, C. grandidieri and C. creolei , and C. creolei is present only in Comoros and the Seychelles. All other Carebara species present in Madagascar have 10 or 11 antennal segments. The main character present in the queens of C. grandidieri and C. voeltzkowi described by Forel, is the presence of nine antennal segments, while the morphological differences between these specimens are not sufficient to merit placement in separate species. It is possible that C. grandidieri represents a species complex, with two or three different species, but much more detailed studies, including molecular studies, will likely be required to confirm this.

The main variations between the queens (Figure 40) of the two species are in the form of the head and the posterodorsal corner of the propodeum. After examining queens collected in different localities we see that the shape of the head is very variable in the queen caste, with some populations with lateral margins of the head convex, and others nearly straight. In the queen caste of C. grandidieri , the posterodorsal corner of the propodeum is unarmed, while in C. voeltzkowi it has a pair of small and angulate triangular teeth, though we found that this character is highly variable in the queens and the major worker caste. For this reason, we are including C. voeltzkowi as a junior synonym of C. grandidieri . In general, the queen of C. grandidieri can be separated from others by the following combination of characters: head longer than wide, in full-face view nearly rectangular, or nearly subquadrate, posterior margin of head slightly concave in the middle, posterolateral corners rounded; lateral margins straight to slightly convex; mandible with six or seven teeth; anterior margin of clypeus slightly concave; antennae with nine segments; ocelli present, eyes present.

Material examined.

COMOROS: Anjouan, -12.25764, 44.38915, 20 m, along roadside, 27.i.2009, (B.L. Fisher et al.); Mohéli Ouallah, -12.32717, 43.65952, 10 m, coastal scrub, 17.i.2009, (B.L. Fisher et al.); Mohéli Ouallah, -12.30668, 43.66407, 275 m, rainforest, 21-24.i.2009, (B.L. Fisher et al.); Mohéli Lac Boundouni, -12.37915, 43.85165, 25 m, dry forest, 20-21.i.2009, (B.L. Fisher et al.); Mohéli Ouallah, -12.30353, 43.66827, 500 m, rainforest, 18-22.i.2009, (B.L. Fisher et al.); Mohéli Ouallah, -12.30353, 43.66827, 500 m, rainforest, 18-19.i.2009, (B.L. Fisher et al.). MADAGASCAR: Antananarivo . Réserve Spéciale d’Ambohitantely, Forêt d’ Ambohitantely, 20.9 km 72° NE d’ Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest, 17-22.iv.2001, (Fisher, Griswold et al.); Antananarivo , [ Antananarivo ], ( Camboué); Antananarivo , Ankalalahana, -19.00659, 47.1122, 1375 m, Uapaca Woodland, 29-31.iii.2011, (B.L. Fisher et al.); Antsiranana . Forêt de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest, 1.xii.2003, (B.L. Fisher); Antsiranana , Forêt d’ Antsahabe, 11.4 km 275° W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest, 12.xii.2003, (B.L. Fisher); Antsiranana , Forêt d’ Andavakoera, 21.4 km 75° ENE Ambilobe; 4.6 km 356° N Betsiaka, -13.11833, 49.23, 425 m, rainforest, 15.xii.2003, (B.L. Fisher); Antsiranana , Forêt de Bekaraoka, 6.8 km 60° ENE Daraina, -13.16667, 49.71, 150 m, tropical dry forest, 7.xii.2003, (B.L. Fisher); Antsiranana , Réserve Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest, 3.xii.2004, (B.L. Fisher); Antsiranana , Réserve Analamerana, 28.4 km 99° Anivorano-Nord, -12.74667, 49.49483, 60 m, tropical dry forest, 5.xii.2004, (B.L. Fisher); Antsiranana , Forêt Ambato, 26.6 km 33° Ambanja, -13.4645, 48.55167, 150 m, rainforest, 8.xii.2004, (B.L. Fisher); Antsiranana , Ambondrobe, 41.1 km 175° Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest, 29.xi.2004, (B.L. Fisher); Antsiranana , Forêt Ambato, 26.6 km 33° Ambanja, -13.4645, 48.55167, 150 m, rainforest, 10.xii.2004, (B.L. Fisher); Antsiranana , Ambondrobe, 41.1 km 175° Vohemar, -13.71533, 50.10167, 10 m, littoral rainforest, 30.xi.2004, (B.L. Fisher); Antsiranana , Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, -14.43667, 49.775, 450 m, rainforest, 23.xi.2004, (B.L. Fisher); Antsiranana , Forêt d’Ampombofofo, -12.09949, 49.33874, 25 m, littoral forest, 21-22.xi.2007, (B.L. Fisher et al.); Antsiranana , Nosy Faly, -13.36435, 48.49137, 40 m, open secondary vegetation, 25.ii.2013, (B.L. Fisher et al.); Antsiranana , Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest, 19-24.iii.2001, (Fisher, Griswold et al.); Antsiranana , Réserve Spéciale de l’Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest, 10-16.ii.2001, (Fisher, Griswold et al.); Antsiranana , Forêt d’Anabohazo, 21.6 km 247° WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest, 11-16.iii.2001, (Fisher, Griswold et al.); Antsiranana , Réserve Spéciale de l’Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest, 16-21.ii.2001, (Fisher, Griswold et al.); Antsiranana , Réserve Spéciale de l’Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889, 49.10983, 80 m, tropical dry forest on tsingy, 10-16.ii.2001, (Fisher, Griswold et al.); Antsiranana , Ampasindava, Forêt d’Ambilanivy, 3.9 km 181° S Ambaliha, -13.79861, 48.16167, 600 m, rainforest, 4-9.iii.2001, (Fisher, Griswold et al.); Antsiranana , Forêt d’Ampondrabe, 26.3km 10° NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest, 10.xii.2003, (B.L. Fisher); Antsiranana , Réserve Spéciale d’Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest, 26-31.i.2001, (Fisher, Griswold et al.). Fianarantsoa . Parc National d’Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest, 10-13.ii.2003, (Fisher, Griswold et al.); Fianarantsoa , Parc National d’Isalo, 9.1 km 354° N Ranohira, -22.48167, 45.46167, 725 m, gallery forest, 27-31.i.2003, (Fisher, Griswold et al.); Fianarantsoa , Parc National d’Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland, 9-14.ii.2003, (Fisher, Griswold et al.); Fianarantsoa , Réserve Forestière d’Agnalazaha, Mahabo, 42.9 km 215° Farafangana, -23.19383, 47.723, 20 m, littoral rainforest, 19.iv.2006, (B.L. Fisher et al.); Fianarantsoa , Réserve Spéciale Manombo 24.5 km 228° Farafangana, -23.01583, 47.719, 30 m, rainforest, 20.iv.2006, (B.L. Fisher et al.); Fianarantsoa , Forêt d’Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, grassland, 22-26.i.2003, (Fisher, Griswold et al.). Mahajanga Parc National tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on tsingy, 16-20.xi.2001, (Fisher, Griswold et al.); Mahajanga , Mahavavy River, 6.2 km 145° SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest, 1-5.xii.2002, (Fisher, Griswold et al.); Mahajanga , Parc National de Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest, 8-12.xi.2002, (Fisher, Griswold et al.); Mahajanga , Réserve Spéciale de Bemarivo, 23.8 km 223° SW Besalampy, -16.925, 44.36833, 30 m, tropical dry forest, 19-23.xi.2002, (Fisher, Griswold et al.); Mahajanga , Parc National de Namoroka, 9.8 km 300° WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest, 4-8.xi.2002, (Fisher, Griswold et al.); Mahajanga , Parc National de Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest, 12-16.xi.2002, (Fisher, Griswold et al.); Mahajanga , Forêt de Tsimembo, 8.7 km 336° NNW Soatana, -19.02139, 44.44067, 20 m, tropical dry forest, 21-25.xi.2001, (Fisher, Griswold et al.); Mahajanga , Parc National tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River, -19.13222, 44.81467, 100 m, tropical dry forest on tsingy, 11-15.xi.2001, (Fisher, Griswold et al.); Mahajanga , Réserve Forestière Beanka, 50.7 km E Maintirano, -17.88021, 44.46877, 140 m, tropical dry forest on tsingy, 29.x.-1.xi.2009, (B.L. Fisher et al.); Mahajanga , Réserve Forestière Beanka, 50.2 km E Maintirano, -18.02649, 44.05051, 250 m, tropical dry forest on tsingy, 19-22.x.2009, (B.L. Fisher et al.); Mahajanga , Réserve Forestière Beanka, 52.7 km E Maintirano, -18.0622, 44.52587, 300 m, tropical dry forest on tsingy, 24-27.x.2009, (B.L. Fisher et al.); Mahajanga , Parc National d’Ankarafantsika, Forêt de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest, 2-8.iv.2001, (Fisher, Griswold et al.); Mahajanga , Parc National tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest, 6-10.xi.2001, (Fisher, Griswold et al.); Mahajanga , Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 40 km 306° NW Andranofasika, -16.32083, 46.81067, 130 m, tropical dry forest, 26.iii.-1.iv.2001, (Fisher, Griswold et al.); Mahajanga , Parc National d’Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331° NW Andranofasika, -16.29889, 46.813, 70 m, tropical dry forest, 26.iii.-1.iv.2001, (Fisher, Griswold et al.); Mahajanga , Réserve d’Ankoririka, 10.6 km 13° NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest, 9-14.iv.2001, (Fisher, Griswold et al.); Mahajanga , Forêt de Tsimembo, 11.0 km 346° NNW Soatana, -18.99528, 44.4435, 50 m, tropical dry forest, 21-25.xi.2001, (Fisher, Griswold et al.). Toamasina. Parcelle K7 Tampolo, -17.28333, 49.41667, 10 m, littoral forest, 20.iv.2004, (Malagasy ant team); Toamasina, Forêt d’Analava Mandrisy, 5.9 km 195 º Antanambe, -16.48567, 49.847, 10 m, littoral rainforest, 13.xi.2005, (B.L. Fisher et al.); Toamasina, Mahavelona (Foulpointe), -17.66667, 49.5, 20.v.1993, (A. Pauly); Toamasina, Station Forestière Analamazaotra, Analamazaotra 1.3 km S Andasibe, -18.38466, 48.41271, 980 m, montane rainforest, 11-13.xii.2007, (B.L. Fisher et al.); Toamasina, Réserve Forestière Tampolo, 95.2 km N Toamasina, -17.27808, 49.42853, 20 m, littoral rainforest, 7.v.2008, (B.L. Fisher et al.); Toamasina, Tamatave, (Voeltzkow); Toliara . Parc National de Kirindy Mite, 16.3 km 127° SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest, 6-10.xii.2001, (Fisher, Griswold et al.); Toliara , Réserve Privé Berenty, Forêt de Malaza, Mandraré River, 8.6 km 314° NW Amboasary, -25.00778, 46.306, 40 m, gallery forest, 6.ii.2002, (Fisher, Griswold et al.); Toliara , Forêt de Mite, 20.7 km 29° WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest, 27.ii.-3.iii.2002, (Fisher, Griswold et al.); Toliara , Réserve Privé Berenty, Forêt de Bealoka, Mandraré River, 14.6 km 329° NNW Amboasary, -24.95694, 46.2715, 35 m, gallery forest, 3-8.ii.2002, (Fisher, Griswold et al.); Toliara , Forêt de Mahavelo, Isantoria River, -24.75833, 46.15717, 110 m, spiny forest/thicket, 28.i.-1.ii.2002, (Fisher, Griswold et al.); Toliara , Réserve Spéciale de Cap Sainte Marie, 12.3 km 262° W Marovato, -25.58167, 45.16833, 200 m, spiny forest/thicket, 11-15.ii.2002, (Fisher, Griswold et al.); Toliara , Parc National d’Andohahela, Forêt de Manantalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, -24.81694, 46.61, 150 m, spiny forest/thicket, 12-16.i.2002, (Fisher, Griswold et al.); Toliara , Parc National de Tsimanampetsotsa, 6.7 km 130° SE Efoetse, 23.0 km 175° S Beheloka, -24.10056, 43.76, 25 m, spiny forest/thicket, 18-22.iii.2002, (Fisher, Griswold et al.); Toliara , Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, -18.26667, 45.40667, 1050 m, montane rainforest, 13-17.i.2003, (Fisher, Griswold et al.); Toliara , Réserve Spéciale d’Ambohijanahary, Forêt d’Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala, -18.26, 45.41833, 1100 m, montane rainforest, 16.i.2003, (Fisher, Griswold et al.); Toliara , Forêt de Kirindy, 15.5 km 64° ENE Marofandilia, -20.045, 44.66222, 100 m, tropical dry forest, 28.xi.-3.xii.2001, (Fisher, Griswold et al.); Toliara , Antafoky, -23.48778, 44.0775, 60 m, gallery forest, 9.ii.2002, (Frontier Project); Toliara , Manombo, -22.8123, 43.73932, 165 m, gallery forest, TS3, 22-24.v.2004, (Frontier Wilderness Project); Toliara , Amboasary, -25.03883, 46.3835, 25 m, urban/garden, 9.xii.2006, (B.L. Fisher et al.); Toliara , Manatantely, 8.9km NW Tolagnaro, -24.9815, 46.92567, 100 m, rainforest, 27.xi.2006, (B.L. Fisher et al.); Toliara , Réserve Spéciale Kalambatritra, Ambinanitelo, -23.45707, 46.4473, 1300 m, grassland, 8.ii.2009, (B.L. Fisher et al.); Toliara , Réserve Spéciale Kalambatritra, Ambinanitelo, -23.45373, 46.45773, 1345 m, grassland, 11.ii.2009, (B.L. Fisher et al.); Toliara , Réserve Spéciale Kalambatritra, -23.4185, 46.4583, 1365 m, grassland, 8.ii.2009, (B.L. Fisher et al.); Toliara , Beza-Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest, 23.iv.1997, (B.L. Fisher); Toliara , Makay Mts., -21.21985, 45.32396, 500 m, gallery forest on sandy soil, 25.xi.2010, (B.L. Fisher et al.); Toliara , Makay Mts, -21.22284, 45.32477, 490 m, gallery forest on sandy soil, 24.xi.-1.xii.2010, (B.L. Fisher et al.); Toliara , Makay Mts., -21.20978, 45.34184, 525 m, gallery forest on sandy soil, 27.xi-2.xii.2010, (B.L. Fisher et al.); Toliara , Mahafaly Plateau, 6.2 km 74° ENE Itampolo, -24.65361, 43.99667, 80 m, spiny forest/thicket, 21-25.ii.2002, (Fisher, Griswold et al.); Toliara , Réserve Spéciale de Cap Sainte Marie, 14.9 km 261° W Marovato, -25.59444, 45.14683, 160 m, spiny forest/thicket, 13-19.ii.2002, (Fisher, Griswold et al.); MAYOTTE: Dapani, -12.96279, 45.15037, 135 m, rainforest, 2-4.xii.2007, (B.L. Fisher et al.); MAYOTTE: Mont Benara, -12.87585, 45.15672, 425 m, rainforest, 30.xi.-2.xii.2007, (B.L. Fisher et al.); MAYOTTE: Mont Chongui, -12.95903, 45.13411, 380 m, rainforest, 28-30.xi.2007, (B.L. Fisher et al.); MAYOTTE: Mont Combani, -12.80632, 45.15314, 370 m, rainforest, 25-28.xi.2007, (B.L. Fisher et al.).