Pachycondyla wasmannii (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 : 132-136

Pachycondyla wasmannii (Forel)

(Figures 9, 11, 50-51,64)

Bothroponera wasmannii Forel 1887 : 383. Lectotype worker, present designation, Madagascar, Antsiranana, Nosy Be (C. Keller) AntWeb specimen code CASENT0101039 (MHNG) [examined]. Paralectotype workers, with the same data but coded as CASENT0101040, CASENT0101041 (MHNG) [examined]. [Redescription: Forel, 1891: 128, pl. 4, fig. 4; Dalla Torre, 1893: 37. Misspelled as wasmanni : Emery, 1895: 336, Forel, 1897: 188, 196 with description of queen and male castes. Combination in Ponera (Bothroponera) and additional note on queen by Emery, 1899: 267; in Pachycondyla (Bothroponera) : Emery, 1901: 45, 1911: 78; Combination in Bothroponera : Wheeler, 1922: 1008; in Pachycondyla : Bolton, 1995: 311].

Worker diagnosis:

With head in full-face view, antennal scape not surpassing the posterior cephalic margin; dorsum of head and mesosoma not finely striate; basal half of antennal scape rounded; dorsum of mesosoma and petiole node with short and thin erect hairs; junction of propodeal dorsum and declivitous surface at a distinct angle; mesopleural suture distinct; antennal segments 6, 7, and 8 nearly as wide as long; with petiole in dorsal view, the posterior margin straight or with weak median notch; anterior half of fourth abdominal tergite (gastral tergite 2) covered with numerous and very dense small punctures between larger ones, mesosoma and petiole node with moderate such sculpture; outer surface of hind tibia usually with erect hairs; integument matte.

Worker measurements (n=12): HL: 1.96-2.37, HW: 1.72-2.21, CI: 87-94, SL: 1.32-1.78, SI: 76-83, PW: 1.28-1.79, WL: 2.50-3.36, NL: 0.94-1.20, NW: 1.07-1.48, NH: 1.27-1.64, DNI: 105-132, LNI: 124-159.

Description:

Worker. In full-face view, head broadest behind level of eyes, the sides slightly convex throughout their length; the posterior margin feebly concave; eyes quite large, maximum diameter nearly equal to greatest width of antennal scape. Scape subcylindrical, relatively short, not surpassing posterior cephalic margin; segments 6, 7, and 8 almost as long as broad. Anteromedian margin of clypeus truncate and either straight or very weakly emarginated medially. Mandibles triangular, masticatory margins armed with seven to eight teeth and denticles. In profile, dorsal outline of mesosoma without uninterrupted convexity; propodeal dorsum meeting declivitous surface in a distinct angle; mesopleural sulcus distinct and complete. Hind legs with rounded basitarsus. Petiole nodiform and thick, with anterior and posterior margins that in dorsal view are respectively broadly convex and straight with weak median notch. Head finely reticulate-punctate to finely ruguloreticulate, interspersed with large punctures behind level of eyes.

Mandibles striate, with sparse punctures from which hairs arise. Dorsum of mesosoma, petiole node, and first gastral tergite covered with large punctures, the spaces between which are reticulate-punctate. Lateral portion of pronotum, mesopleuron, and lower half of propodeum generally finely reticulate-rugulose or with dense, fine rugulation. Second gastral tergite particularly with very dense, piligerous, small pits between large punctures. Standing erect hairs present on dorsum of the body, with very abundant pubescence; outer surface of hind tibiae covered with erect hairs. Dark brown to black in color with usually lighter appendages. Most specimens fairly matte except those collected from rainforest habitats.

Queen. Measurements (n=7): HL: 2.20-2.39, HW: 2.08-2.24, CI: 92-95, SL: 1.62-1.82, SI: 77-84, EL: 0.39-0.41, OI: 17-19, PW: 1.63-1.87, WL: 3.31-3.71, NL: 0.99-1.16, NW: 1.34-1.48, NH: 1.31-1.51, DNI: 127-148, LNI: 125-136. Winged queens are very similar to workers, but have a body modified with the general characteristics of the queen caste. Winged queen has a much broader head. Ergatoid queens also present in P. wasmannii , which look comparable to worker caste but have one ocellus and reduced thoracic sclerites.

Discussion:

Although P. wasmannii is very similar to P. vazimba , it can be distinguished readily by its larger size, with a distinct and complete mesopleural suture, antennal segments 6, 7, and 8 nearly as wide as long, and more erect hairs on the dorsum of mesosoma and petiole node.

Apparently there are two forms found within this species. These are geographically isolated due to the presence of high mountain chains in northwestern Madagascar. The first form, in which gastral tergite 2 is densely, finely punctate between larger punctures and has a matte integument, occupies the west and southwest regions of Madagascar. In contrast, the second form is characterized by worker specimens that have a shining integument, and gastral tergite 2 is covered with nearly effaced, closely spaced, small punctures between larger, shallow punctures. This form has been collected in the northwest and northeast of the island. However, workers with intermediate degrees of these phenotypic variations also occur.

Distribution and biology:

Pachycondyla wasmannii is an endemic, widespread species of the Malagasy region. It occurs generally in western Madagascar and in Anjouan of the Comoros Islands (Fig. 64). Most frequently this species has been collected from the dry forests and woodland habitats in the west of Madagascar. However, it is known also from subhumid gallery forests in the western slopes, littoral, transitional, and humid forests in the north and southeast of the island, and from disjunct montane rainforests on the high plateau. This species is also capable of colonizing human-modified habitats. There are two morphological forms; the first occurs in the dry and gallery forest habitats in the west of Madagascar, whereas the second is generally known from the transitional mesic forests and some of the rainforest sites in the north and northeast of the island. Across its distribution range, P. wasmannii dominates the Pachycondyla in dry forest habitats where it is sympatric only with P. vazimba . In its remaining distribution range, other species within the genus co-occur with P. wasmannii , and P. cambouei , which is widely distributed throughout the humid forests, becomes the dominant species. Pachycondyla wasmannii nests mainly in rotten logs; however, it can also be found on rotten sticks and branches on the ground, in soil layers, under rocks, beneath litter moss on rocks, and under rotten logs. It typically forages on the forest floor and through leaf litter, and very rarely on low-growing vegetation. Workers of this species usually fake death (thanatosis) after a nest disturbance.

One interesting geographic point about P. wasmannii is its presence on Anjouan Island in the Comoros but not Mayotte Island, although the latter is geographically closer to Madagascar. How it arrived on Comoros could shed light on our understanding of the historical biogeography and the origins of the Malagasy species of Pachycondyla .

Other material examined:

THE COMOROS: Anjouan, Mount Ntringui, -12.19865, 44.41866, 740 m, montane forest (B.L. Fisher et al.) (CASC); Mount Ntringui, -12.19641, 44.41791, 550 m, rainforest (B.L. Fisher et al.) (CASC). MADAGASCAR: Antananarivo: RS Ambohitantely, F d Ambohitantely, 20.9 km 72° NE d Ankazobe, -18.22528, 47.28683, 1410 m, montane rainforest (Fisher, Griswold et al.) (CASC); Tsinjoarivo, E. Ambatolampy, dry forest (W.L. Brown) (MCZC), Antsiranana: [ Nosibé] ( Völtzkow) (MHNG); [Nossi bé p. Madagascar] (C. Keller) (MHNG); 14 km W Cap Est, Ambato, -15.29139, 50.33806, 200 m, secondary raiforest (Alpert et al.) (MCZC); 15 km S Sambava, 10 m, coastal forest (J.M. Betsch) (MCZC); 48 km ENE Morondava, -20.06667, 44.65, tropical dry forest (D.M. Olson) (MCZC); 5 km S Antalaha, Ambodikofo Hill (G.D. Alpert) (MCZC); 5 km SW Antalaha, -14.93806, 50.26167, 50 m, secondary forest (G.D. Alpert) (MCZC); 84 km SW Sambava to Andapa, 70-160 m, degraded forest (W.L. and D.E. Brown) (MCZC); Ambanja, -13.68268, 48.45245, 30 m, urban/garden (B.L. Fisher et al.) (CASC); Ambondrobe, 41.1 km 175° Vohemar, -13.71533, 50.10167, 10 m, littoral 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); Ankarana, -12.9, 49.1, 100 m (G.D. Alpert) (MCZC); F Ambanitaza, 26.1 km 347° Antalaha, -14.67933, 50.18367, 240 m, rainforest (B.L. Fisher) (CASC); F Ambato, 26.6 km 33° Ambanja, - 13.4645, 48.55167, 150 m, rainforest (B.L. Fisher) (CASC); F d' Andavakoera, 21.4 km 75° ENE Ambilobe; 4.6 km 356° N Betsiaka, -13.11833, 49.23, 425 m, rainforest (B.L. Fisher) (CASC); F d' Antsahabe, 11.4 km 275° W Daraina, -13.21167, 49.55667, 550 m, tropical dry forest (B.L. Fisher et al.) (CASC); F d'Ampombofofo, - 12.09949, 49.33874, 25 m, littoral forest (B.L. Fisher et al.) (CASC); F d'Ampondrabe, 26.3 km 10° NNE Daraina, -12.97, 49.7, 175 m, tropical dry forest (B.L. Fisher) (CASC); PN Sahamalaza, F d'Anabohazo, 21.6 km 247° WSW Maromandia, -14.30889, 47.91433, 120 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F d'Analabe, 30.0 km 72° ENE Daraina, -13.08333, 49.90833, 30 m, littoral rainforest (B.L. Fisher) (CASC); F de Bekaraoka, 6.8 km 60° ENE Daraina, -13.16667, 49.71, 150 m, tropical dry forest (B.L. Fisher et al.) (CASC); F de Binara, 7.5 km 230° SW Daraina, -13.255, 49.61667, 375 m, tropical dry forest (B.L. Fisher) (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 d'Orangea, 3.6 km 128° SE Remena, -12.25889, 49.37467, 90 m, littoral rainforest (Fisher, Griswold et al.) CASC; Marojejy RNI #12, -14.44533, 49.78564, 375 m, rainforest (G.D. Alpert) (MCZC); Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez), -12.32278, 49.33817, 180 m, tropical dry forest (Alpert et al.) (CASC); Nosy Be, 4 km ESE Andoany (=Hellville), -13.41667, 48.3, 200 m, rainforest (P.S. Ward) (PSWC); Nosy Be, Lokobe Forest, - 13.41639, 48.30722, 20 m (G.D. Alpert) (MCZC); Nosy Be, RNI Lokobe, 6.3 km 112° ESE Hellville, -13.41933, 48.33117, 30 m, rainforest (Fisher, Griswold 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) (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, -12.50035, 49.175, 885 m, montane rainforest (B.L. Fisher 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, Pic Bades, -12.5186, 49.18625, 900 m, montane rainforest (B.L. Fisher et al.) (CASC); RS Manongarivo, 10.8 km 229° SW Antanambao, -13.96167, 48.43333, 400 m, rainforest (B.L. Fisher) (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); RS Ankarana, 7 km SE Matsaborimanga, -12.9, 49.11667, 150 m, rainforest (P.S. Ward) (MCZC); RS Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467, 49.37383, 225 m, tropical dry forest (B.L. Fisher) (CASC); RS Analamerana, 28.4 km 99° Anivorano-Nord, -12.74667, 49.49483, 60 m, tropical dry forest (B.L. Fisher) (CASC); RS Ankarana, -12.90056, 49.14722, 150 m, dry forest (Alpert et al.) (MCZC); RS d'Ambre, 3.5 km 235° SW Sakaramy, -12.46889, 49.24217, 325 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ankarana, 13.6 km 192° SSW Anivorano Nord, -12.86361, 49.22583, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Ankarana, 22.9 km 224° SW Anivorano Nord, -12.90889,49.10983, 80 m, tropical dry forest (Alpert et al.) (CASC); PN Marojejy, 8 km NW Manantenina, -14.43333, 49.78333, 450 m, montane rainforest (E.L. Quinter) (CASC); Vohemar, -13.37723, 50.0205, 25 m, cultivated land (B.L. Fisher et al.) (CASC); Makirovana Forest: -14.1604, 49.9522, 550 m, rainforest (B.L. Fisher et al.) (CASC); Makirovana Forest, - 14.17066, 49.95409, 415 m and 225 m, rainforest (B.L. Fisher et al.) (CASC); Makirovana Forest, -14.10295, 50.01984, 390 m, rainforest (B.L. Fisher et al.) (CASC); Fianarantsoa: 15 km E Sakaraha, -22.9, 44.68333, 760 m, tropical dry forest (P.S. Ward) (MCZC); 30 km NNW Ranohira, Isalo Nat. Park, -22.31722, 45.29333, 455 m, canopy forest (G.D. Alpert) (MCZC); F d'Analalava, 29.6 km 280° W Ranohira, -22.59167, 45.12833, 700 m, tropical dry forest (Fisher, Griswold et al.) (CASC); F d'Atsirakambiaty, 7.6 km 285° WNW Itremo, -20.59333, 46.56333, 1550 m, montane rainforest (Fisher, Griswold et al.) (CASC); PN Isalo, Ranohira, Canyon de Sinze, - 22.48333, 45.55, 800 m, rainforest (E. Rajeriarison) (MCZC); PN Ranomafana, -21.01667, 47.01667, 700 m, montane forest (E. Rajeriarison) (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 Isalo, 9.1 km 354° N Ranohira, - 22.48167, 45.46167, 725 m, gallery forest (Fisher, Griswold et al.) CASC; PN Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, -22.29833, 45.35167, 990 m, Uapaca woodland (Fisher, Griswold et al.) (CASC); PN Isalo, Sahanafa River, 29.2 km 351° N Ranohira, -22.31333, 45.29167, 500 m, gallery forest (Fisher, Griswold et al.) (CASC); 9 km NNW Ranohira, PN Isalo, -22.48333, 45.38333, 800 m, rainforest (P.S. Ward) (MCZC); Tsaranoro, 32.8 km 229° Ambalavao, -22.08483, 46.77633, 950 m, rainforest (B.L. Fisher et al.) (CASC); Mahajanga: 124 km SE Mahajanga (G.D. Alpert) (MCZC); Ampamakiambato, 45 km SW Ambanja, -13.97545, 48.15929, 145 m, disturbed forest in tsingy (B.L. Fisher et al.) (CASC); F Ambohimanga, 26.1 km 314° Mampikony, -15.96267, 47.43817, 250 m, tropical dry forest (B.L. Fisher) (CASC); F de Tsimembo, 11.0 km 346° NNW Soatana, - 18.99528, 44.4435, 50 m, tropical dry forest (Fisher, Griswold Arthropod Team) (CASC); F de Tsimembo, 8.7 km 336° NNW Soatana, -19.02139, 44.44067, 20 m, tropical dry forest (Fisher, Griswold Arthropod Team) (CASC); Mahavavy River, 6.2 km 145° SE Mitsinjo, -16.05167, 45.90833, 20 m, gallery forest (Fisher, Griswold et al.) (CASC); Manerinerina, 76.6 km N Antsohihy, -14.10744, 48.11046, 247 m, disturbed forest (B.L. Fisher et al.) (CASC); PN Ankarafantsika, SF Ampijoroa, 5.4 km 331° NW Andranofasika, -16.29889, 46.813, 70 m, tropical dry forest (Fisher, Griswold et al.) CASC; PN Ankarafantsika, Ampijoroa SF, 40 km 306° NW Andranofasika, - 16.32083, 46.81067, 130 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Ankarafantsika, F de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, -16.22806, 47.14361, 135 m, tropical dry forest (Fisher, Griswold et al.) CASC; PN Baie de Baly, 12.4 km 337° NNW Soalala, -16.01, 45.265, 10 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 16.9 km 317° NW Vilanandro, -16.40667, 45.31, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 17.8 km 329° WNW Vilanandro, -16.37667, 45.32667, 100 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Namoroka, 9.8 km 300° WNW Vilanandro, -16.46667, 45.35, 140 m, tropical dry forest (Fisher, Griswold et al.) (CASC); PN Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, -18.70944, 44.71817, 150 m, tropical dry forest on tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River,-19.13222, 44.81467, 100 m, tropical dry forest on tsingy (Fisher-Griswold Arthropod Team) (CASC); PN Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, -19.14194, 44.828, 50 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); Réserve d'Ankoririka, 10.6 km 13° NE de Tsaramandroso, -16.26722, 47.04861, 210 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RF Beanka, 50.2 km E Maintirano, -18.02649, 44.05051, 250 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 52.7 km E Maintirano, -18.0622, 44.52587, 300 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 53.6 km E Maintirano, -18.04014, 44.53394, 272 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RF Beanka, 54.3 km E Maintirano, -18.06009, 44.54086, 262 m, tropical dry forest on tsingy (B.L. Fisher et al.) (CASC); RS Bemarivo, 23.8 km 223° SW Besalampy, - 16.925, 44.36833, 30 m, tropical dry forest (Fisher, Griswold et al.) (CASC); RS Marotandrano, Marotandrano 48.3 km S Mandritsara, -16.28322, 48.81443, 865 m, transitional humid forest (B.L. Fisher et al.) (CASC); Toamasina: 1 km W Andampibe, Cap Masoala, -15.69361, 50.18139, 125 m, rainforest (G.D. Alpert) (MCZC); 8 km SW Cap Est, -15.30694, 50.45417, 15 m, littoral forest (G.D. Alpert) (MCZC); Toliara: 29 km NNW Ranohira, PN Isalo, -22.31614, 45.29625, 490 m, canopy forest (G.D. Alpert) (MCZC); 48 km ENE Morondava, Kirindy, - 20.06667, 44.65, 30 m, tropical dry forest (D.M. Olson) (CASC); 48 km ENE Morondava, Kirindy Forest, - 20.07444, 44.67611, 100 m, tropical dry forest (P. Rabeson) (MCZC); 50 kms N Morondava, -20.06667, 44.58333, primary dry forest (A. Pauly) (CASC); 9 km NNW Ranohira, P.N. Isalo, -22.48333, 45.38333, 800 m, rainforest (P.S. Ward) (BMNH); Andohahela, Parcel #1 versante E, 300 m, rainforest (L. Bartolozzi, S. Tiati & C. Raharimina) (MCZC); Bekonazy = 0.5 km S (n. of Morondava), forest with baobab (W.L. Brown) (MCZC); Beza Mahafaly, 27 km E Betioky, -23.65, 44.63333, 135 m, tropical dry forest (B.L. Fisher) (CASC); Fiherenana, - 23.17694, 43.96083, 100 m, gallery forest (Frontier Project) (CASC); 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 de Beroboka, 5.9 km 131° SE Ankidranoka, -22.23306, 43.36633, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); F de Kirindy, 15.5 km 64° ENE Marofandilia, -20.06855, 44.65956667, 30 m, tropical dry forest (B.L. Fisher) (CASC); F de Mite, 20.7 km 29° WNW Tongobory, -23.52417, 44.12133, 75 m, gallery forest (Fisher-Griswold Arthropod Team) (CASC); F de Petriky, 12.5 km W 272° Tolagnaro, -25.06167, 46.87, 10 m, littoral rainforest (B.L. Fisher) (CASC); F de Zombitse near Sakaraha, 650 m (E.S. Ross) (MCZC); F Ivohibe 55.0 km N Tolagnaro, -24.569, 47.204, 200 m, rainforest (B.L. Fisher et al.) (CASC); F Vohidava 88.9 km N Amboasary, -24.24067, 46.28783, 500 m, spiny forest/dry forest transition (B.L. Fisher et al.) (CASC); Makay Mts., -21.227, 45.33222, 475 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.22336, 45.32628, 480 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.22344, 45.3135, 550 m, gallery forest with bamboo (B.L. Fisher et al.) (CASC); Makay Mts., -21.31664, 45.1296, 620 m, dry forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.21836, 45.3106, 510 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.20978, 45.34184, 525 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.30997, 45.12946, 590 m, dry forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., -21.21761, 45.33917, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Makay Mts., - 21.23343, 45.32913, 460 m, gallery forest with bamboo (B.L. Fisher et al.) CASC); Makay Mts., -21.21985, 45.32396, 500 m, gallery forest on sandy soil (B.L. Fisher et al.) (CASC); Manatantely, 8.5 km NW Tolagnaro, - 24.9875, 46.92617, 85 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); Manombo, -22.8123, 43.73932, 165 m, gallery forest, TS3 (Frontier Wilderness Project) (CASC); Manombo, -22.80257, 43.76505, 227 m, dry forest, SC2 (Frontier Wilderness Project) (CASC); Manombo, -22.8123, 43.73932, 165 m, gallery forest, TS3 (Frontier Wilderness Project) (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 Kirindy Mite, 16.3 km 127° SE Belo sur Mer, -20.79528, 44.147, 80 m, tropical dry forest (Fisher-Griswold Arthropod Team) (CASC); PN Zombitse, 19.8 km 84° E Sakaraha, -22.84333, 44.71, 770 m, tropical dry forest (Fisher, Griswold et al.) (CASC); Réserve Beza Mahafaly, Parcel 1, -23.65833, 44.62889, 185 m (P. Rabeson) (MCZC); RP Berenty, - 25.01667, 46.3, 25 m, tropical dry forest (B.L. Fisher) (CASC); RP Berenty, F de Bealoka, Mandraré River, 14.6 km 329° NNW Amboasary, -24.95694, 46.2715, 35 m, gallery forest (Fisher-Griswold Arthropod Team) (CASC); RP Berenty, F de Malaza, Mandraré River, 8.6 km 314° NW Amboasary, -25.00778, 46.306, 40 m, gallery forest (Fisher-Griswold Arthropod Team) (CASC); RP Berenty, -25.00667, 46.30333, 85 m, gallery forest (Rin'Ha, Irwin) (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); Sept Lacs, -23.52833, 44.15556, 80 m, gallery forest (Frontier Project) (CASC); Southern Isoky-Vohimena Forest, 59 km NE Sakaraha, -22.46667, 44.85, 730 m, tropical dry forest (B.L. Fisher) (CASC); Vohibasia Forest, 59 km NE Sakaraha, -22.46667, 44.85, 780 m, tropical dry forest (B.L. Fisher) (CASC); Zombitsy Forest, 15 km NE Sakaraha, -22.8925, 44.67833, 750 m, secondary forest (E. Rajeriarison) (MCZC).