Taxonomic revision of the genus Prionopelta (Hymenoptera, Formicidae) in the Malagasy region Author Overson, Rick Author Fisher, Brian L. text ZooKeys 2015 507 115 150 http://dx.doi.org/10.3897/zookeys.507.9303 journal article http://dx.doi.org/10.3897/zookeys.507.9303 1313-2970-507-115 83353FFD8729496E92402EFA8FC86BCE 83353FFD8729496E92402EFA8FC86BCE Taxon classification Plantae Hymenoptera Formicidae Prionopelta descarpentriesi Santschi Figs 1C, D, 3D, 4B, C, E, F, 5, 6 Prionopelta descarpentriesi Santschi, 1924b:195. Type material. Lectotype, pinned worker, CASENT0101548 [designated here], MADAGASCAR, Ikelivia, 30.ix.1923 ( Descarpentries ) (NHMB) [examined]. Paralectotypes, one pinned worker CASENT0101547 with same data as lectotype (NHMB) [examined]. Diagnosis. Prionopelta descarpentriesi can be identified by the following combination of characters: twelve antennal segments; median cephalic band lacking a thin suture that is swollen above the surrounding integument; placement of cephalic foveae ranging from sparse to dense, but always at minimum, with at least several clusters of foveae directly adjacent to one another (if nowhere else, then medially in full-face view); if all foveae on the head are directly adjacent so that no flat, shining space is present between foveae, then foveae are large and accompanied by pronotal sculpture which is characterized as being both similar in size to that on the head and not consisting of smaller foveae or punctures; eye appearing as either an asymmetrical dark patch which appears to be a stain in the cuticle that is flush with its surrounding integument, or a single, slightly rounded glob with no definable subunits. Worker measurements (N=25). HL 0.42-0.53 (0.48); HW 0.32-0.43 (0.38); SL 0.22-0.3 (0.26); WL 0.46-0.6 (0.52); PetL 0.11-0.2 (0.16); PetW 0.17-0.26 (0.22); T1W 0.29-0.37 (0.33); CI 74.64-85.1 (79.6); PI 118.23-173.29 (141.94); SI 61.46-72.38 (67.31). Worker description. Posterior margin of the head straight to weakly concave in full-face view; spacing of cephalic foveae highly variable, ranging from individuals with dense, directly adjacent foveae covering the entire head (known only from far eastern and northern Madagascar, see morphotype descriptions below), to individuals with foveae more widely spaced so that shining areas are visible between; median cephalic band devoid of foveae ranging from wide to extremely narrow but never appearing as a linear suture that is uniformly swollen above the level of the surrounding integument; apical tooth intermediate in length; evenly-space pronotal foveae range from shallow to deep; shallow foveae present on mesonotum and propodeum. Distribution and ecology. This widespread species has been collected from leaf litter from 10-1860 meters of elevation. While found most commonly in rainforest and montane rainforest, it has also been collected in Uapaca woodland, littoral rainforest, and tropical dry forest (Fig. 13). Prionopelta descarpentriesi has been collected in forest litter, under moss, rocks, and logs, as well as inside rotten logs and underground in soil. Taxonomic notes. Three generalized morphotypes can be distinguished within this taxon; these vary in density of cephalic foveae and other co-occurring traits (Fig. 6). In morphotype A, the majority of foveae are equidistantly spaced and separated by a span of shining integument of around one foveal diameter (Figs 1C, 5A, 6A). These foveae appear cleanly scooped from the surface of the integument and largely lack raised margins. Morphotype B, which is intermediate between A and C, has denser cephalic foveae covering almost the entire head, however foveae are smaller and more delicate than those of morphotype C and raised ridges between foveae are less pronounced (Fig. 6B). Pronotal sculpture in morphotype B usually consists of several sizes of foveae along with some punctures. Morphotype C possesses large, dense cephalic foveae that cover the entire head, accompanied by a pronounced network of raised, jagged ridges between foveae (Figs 1D, 3D, 4C, 6C). Morphotype C additionally has large, deep, and evenly spaced pronotal foveae that are similar in size to those on the head, and this morphotype lacks smaller foveae or punctures on the pronotum. Width of the median cephalic band devoid of foveae is widest in morphotype A and narrowest in morphotype C. Figure 6. A, B, C Head in full-face view displaying variation in Prionopelta descarpentriesi for cephalic sculpture. A (CASENT0034837) "morphotype-A" B (CASENT0480792) "morphotype-B" C (CASENT0191895) "morphotype-C" . Prionopelta descarpentriesi is much more morphologically variable than the other species treated in this revision and very possibly represents a species complex. This could explain why Prionopelta descarpentriesi is abundant, geographically widespread, and morphologically variable. Morphotype C, which is restricted to a band bordering the coast of eastern and northern Madagascar (Fig. 14), is recognizably distinct from the majority of individuals from interior populations of the species. At several localities where morphotype C is present, individuals from morphotype A and B are also present with little to no evidence of character blending between morphotypes: Galako, Makirovana, Morojejy Nature Reserve, and Sahafina. However, at other locales such as Ambohijanahary, Montagne d'Ambre , and Vohemar, a bewildering array of intermediate forms have been collected, blurring the lines between the three morphotypes. Collecting nest series from the aforementioned locations to determine whether this population-level variation is intra- or intercolonial is an important first step in ultimately understanding how morphological variation is partioned in this taxon as currently dilineated. Under the above scheme, the lectotype and paralectotype of Prionopelta descarpentriesi would be considered morphotype A. Jules Descarpentries (1881-1927) collected the types for this species on 30 September 1923 and the collection locale was subsequently noted as "Ikelivia" , making it difficult to determine a more precise location based on modern place names. Historical records indicate that Descarpentries resided in Tulear, and worked as a topographic surveyor and entomologist. According to records, he was active around Tsaratanana and Fianarantsoa (specifically Andringitra) and often traveled with H. Perrier de la Bathie . Given that morphotype A is reasonably widespread across Madagascar, including Tsaratanana and Andringitra, it is possible that the type specimens from "Ikelivia" were collected in either of these localities. To further complicate matters, most of Descarpentries's specimens, which were destined to the Paris Museum, were sold on the side by a member of the staff. Before the specimens were sold, the labels were changed to hide their true ordinance ( Jeannel 1951 , ( Anonymous 1970 )). Non-type material. MADAGASCAR: Antananarivo, Ankalalahana, 19.0066°S , 47.1122°E , 1375 m, Uapaca woodland, 29.iii.2011 ( B .L.Fisher et al.); Antananarivo, Foret de galerie, Andranorovitra, 24.0 km NNE Ankazobe, 18.1124°S , 47.1976°E , 1491 m, disturbed gallery montane forest, 2.vi.2008 (B.L.Fisher et al.); Antananarivo, Navoatra I Non Protected Area, 7.64 km NW Arivonimamo, 18.9781°S , 47.1193°E , 1373 m, Uapaca woodland, 6.v.2010 (Andrianjaka Ravelomanana); Antananarivo, Reserve Naturelle Sohisika, Sohisika 24.6 km NNE Ankazobe, 18.1032°S , 47.1869°E , 1464 m, gallery montane forest, 1.vi.2008 (B.L.Fisher et al.); Antananarivo, Reserve Speciale d'Ambohitantely , 18.2244°S , 47.2774°E , 1490 m, montane forest, 9.iii.2012 (B.L.Fisher et al.); Antananarivo, Reserve Speciale d'Ambohitantely , Foret d'Ambohitantely , 20.9 km 72° NE d'Ankazobe , 18.2253°S , 47.2868°E , 1410 m, montane rainforest, 17.iv.2001 (Fisher, Griswold et al.); Antananarivo, Reserve Speciale d'Ambohitantely , Foret d'Ambohitantely , Jardin Botanique, 24.1 km 59° NE d'Ankazobe , 18.1714°S , 47.2818°E , 1620 m, montane rainforest, 17.iv.2001 (Fisher, Griswold et al.); Antananarivo, Tsimbazaza, 18.928°S , 47.527°E , 1300 m, park/garden, 16.xii.2006 (B.L.Fisher et al.); Antsiranana, 6.5 km SSW Befingotra, Res . Anjanaharibe-Sud, 14.75°S , 49.5°E , 875 m, rainforest, 19.x.1994 (B.L.Fisher); Antsiranana, 9.2 km WSW Befingotra, Res . Anjanaharibe-Sud, 14.75°S , 49.4667°E , 1280 m, montane rainforest, 5.xi.1994 (B.L.Fisher); Antsiranana, Ambondrobe, 41.1 km 175° Vohemar, 13.7153°S , 50.1017°E , 10 m, littoral rainforest, 29.xi.2004 (B.L.Fisher); Antsiranana, Ampasindava, Andranomatavy Forest, 13.669°S , 47.9877°E , 149 m, disturbed dry forest, 6.x.2013 ( B .L.Fisher et al.); Antsiranana, Ampasindava, Andranomatavy Forest, 13.663°S , 47.9794°E , 543 m, disturbed dry forest, 6.x.2013 (B.L.Fisher et al.); Antsiranana, Binara Forest, 13.2621°S , 49.6067°E , 559 m, degraded rainforest, 18.x.2013 (B.L.Fisher et al.); Antsiranana, Binara Forest, 13.2621°S , 49.605°E , 692 m, rainforest, 18.x.2013 (B.L.Fisher et al.); Antsiranana, Binara Forest, 13.2639°S , 49.5992°E , 1065 m, rainforest, 18.x.2013 (B.L.Fisher et al.); Antsiranana, Foret Ambanitaza, 26.1 km 347° Antalaha, 14.6793°S , 50.1837°E , 240 m, rainforest, 26.xi.2004 (B.L.Fisher); Antsiranana, Foret d' Andavakoera, 21.4 km 75° ENE Ambilobe; 4.6 km 356° N Betsiaka, 13.1183°S , 49.23°E , 425 m, rainforest, 15.xii.2003 (B.L.Fisher); Antsiranana, Foret d' Antsahabe, 11.4 km 275° W Daraina, 13.2117°S , 49.5567°E , 550 m, tropical dry forest, 12.xii.2003 (B.L.Fisher); Antsiranana, Foret d'Ampondrabe , 26.3 km 10° NNE Daraina, 12.97°S , 49.7°E , 175 m, tropical dry forest, 10.xii.2003 (B.L.Fisher); Antsiranana, Foret de Bekaraoka, 6.8 km 60° ENE Daraina, 13.1667°S , 49.71°E , 150 m, tropical dry forest, 7.xii.2003 (B.L.Fisher); Antsiranana, Foret de Binara, 7.5 km 230° SW Daraina, 13.255°S , 49.6167°E , 375 m, tropical dry forest, 1.xii.2003 (B.L.Fisher); Antsiranana, Foret de Binara, 9.1 km 233° SW Daraina, 13.2633°S , 49.6033°E , 800 m, rainforest, 3.xii.2003 (B.L.Fisher); Antsiranana, Foret de Binara, 9.4 km 235° SW Daraina, 13.2633°S , 49.6°E , 1100 m, montane rainforest, 5.xii.2003 (B.L.Fisher); Antsiranana, Galoko chain, Mont Galoko, 13.5936°S , 48.7316°E , 1100 m, montane forest, 22.ii.2013 (B.L.Fisher et al.); Antsiranana, Galoko chain, Mont Galoko, 13.5888°S , 48.7286°E , 980 m, montane forest, 22.ii.2013 (B.L.Fisher et al.); Antsiranana, Galoko chain, Mont Galoko, 13.5849°S , 48.7182°E , 520 m, rainforest, 16.ii.2013 (B.L.Fisher et al.); Antsiranana, Galoko chain, Mont Kalabenono, 13.6418°S , 48.6728°E , 643 m, rainforest, 10.x.2013 ( B .L.Fisher et al.); Antsiranana, Galoko chain, Mont Kalabenono, 13.64°S , 48.6737°E , 498 m, rainforest, 10.x.2013 (B.L.Fisher et al.); Antsiranana, Galoko chain, Mont Kalabenono, 13.6461°S , 48.6773°E , 937 m, rainforest, 10.x.2013 (B.L.Fisher et al.); Antsiranana, Makirovana forest, 14.1604°S , 49.9522°E , 550 m, rainforest, 1.v.2011 (B.L.Fisher et al.); Antsiranana, Makirovana forest, 14.1651°S , 49.9477°E , 900 m, montane rainforest, 30.iv.2011 (B.L.Fisher et al.); Antsiranana, Makirovana forest, 14.1667°S , 49.95°E , 715 m, rainforest, 1.v.2011 (B.L.Fisher et al.); Antsiranana, Makirovana forest, 14.1707°S , 49.9541°E , 415 m, rainforest, 28.iv.2011 (B.L.Fisher et al.); Antsiranana, Parc National de Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, 14.4433°S , 49.7433°E , 1325 m, montane rainforest, 18.xi.2003 (B.L.Fisher); Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S , 49.76°E , 775 m, rainforest, 15.xi.2003 (B.L.Fisher et al.); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.4367°S , 49.775°E , 450 m, rainforest, 12.xi.2003 (B.L.Fisher et al.); Antsiranana, Parc National Montagne d'Ambre , 12.2 km 211° SSW Joffreville, 12.5964°S , 49.1595°E , 1300 m, montane rainforest, 2.ii.2001 (Fisher, Griswold et al.); Antsiranana, Parc National Montagne d'Ambre , 3.6 km 235° SW Joffreville, 12.5344°S , 49.1795°E , 925 m, montane rainforest, 20.i.2001 (Fisher, Griswold et al.); Antsiranana, Prov.Antsiranana R.S. Manongarivo 17.3 km 218° SW Antanambao, 14.0217°S , 48.4183°E , 1580 m, montane rainforest, 27.x.1998 (B.L.Fisher); Antsiranana, R.S. Manongarivo, 10.8 km 229° SW Antanambao, 13.9617°S , 48.4333°E , 400 m, rainforest, 8.xi.1998 (B.L.Fisher); Antsiranana, R.S. Manongarivo, 12.8 km 228° SW Antanambao, 13.9767°S , 48.4233°E , 780 m, rainforest, 11.x.1998 (B.L.Fisher); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Antanambao, 13.9983°S , 48.4283°E , 1175 m, montane rainforest, 20.x.1998 (B.L.Fisher); Antsiranana, R.S. Manongarivo, 20.4 km 219° SW Antanambao, 14.0467°S , 48.4017°E , 1860 m, montane rainforest, 3.xi.1998 (B.L.Fisher); Fianarantsoa, 28 km. SSW Ambositra, Ankazomivady, 20.775°S , 47.1683°E , 1670 m, grassland, 11.i.1998 (B.L.Fisher); Fianarantsoa, 3 km W Ranomafana, nr. Ifandiana, 21.25°S , 47.4167°E , 950 m, forest, 27.iv.1989 (P.S.Ward); Fianarantsoa, 43 km S Ambalavao, Res . Andringitra, 22.2333°S , 47°E, 825 m, rainforest, 5.x.1993 (B.L.Fisher); Fianarantsoa, 45 km S. Ambalavao, 22.2167°S , 47.0167°E , 785 m, rainforest, 25.ix.1993 (B.L.Fisher); Fianarantsoa, 7.6 km 122 ° Kianjavato, Foret Classee Vatovavy, 21.4°S , 47.94°E , 175 m, rainforest, 6.vi.2005 (B.L.Fisher et al.); Fianarantsoa, 9.0 km NE Ivohibe, 22.4267°S , 46.9383°E , 900 m, rainforest, 12.xi.1997 (B.L.Fisher (Sylvain)); Fianarantsoa, Ampangabe I Non Protected Area, 21.4 km W Itremo, 20.6111°S , 46.6069°E , 1414 m, savannah woodland, 21.iii.2010 (Andrianjaka Ravelomanana); Fianarantsoa, Antapia I Non Protected Area, 26.43 km SW Ambositra, 20.7197°S , 47.0868°E , 1495 m, Uapaca woodland, 3.ii.2010 (Andrianjaka Ravelomanana); Fianarantsoa, Antohatsahomby V Non Protected Area, 22.63 km NW Itremo, 20.5672°S , 46.5792°E , 1726 m, Uapaca woodland, 18.iii.2010 (Andrianjaka Ravelomanana); Fianarantsoa, Foret d'Atsirakambiaty , 7.6 km 285° WNW Itremo, 20.5933°S , 46.5633°E , 1550 m, montane rainforest, 22.i.2003 ( Fisher , Griswold et al.); Fianarantsoa, Mampiarika I Non Protected Area, 28.08 km SW Ambositra, 20.7344°S , 47.0836°E , 1480 m, Uapaca woodland, 31.i.2010 (Andrianjaka Ravelomanana); Fianarantsoa, Mampiarika III Non Protected Area, 28.93 km SW Ambositra, 20.7358°S , 47.084°E , 1487 m, Uapaca woodland, 1.ii.2010 (Andrianjaka Ravelomanana); Fianarantsoa, Parc National Befotaka-Midongy, Papango 27.7 km S Midongy-Sud, Mount Papango, 23.8352°S , 46.9637°E , 940 m, rainforest, 13.xi.2006 (B.L.Fisher et al.); Fianarantsoa, Parc National d'Isalo , 9.1 km 354° N Ranohira, 22.4817°S , 45.4617°E , 725 m, gallery forest, 27.i.2003 (Fisher, Griswold et al.); Fianarantsoa, Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S , 47.4333°E , 1100 m, montane rainforest, 27.iii.2003 (Fisher, Griswold et al.); Fianarantsoa, Parc Nationale Ranomafana: Talatakely, 21.2483°S , 47.4267°E , in guava forest, 9.iv.1998 (CE Griswold, DH Kavanaugh, ND Penny, MJ Raherilalao, JS Ranorianarisoa, J Schwei); Fianarantsoa, R.S. Ivohibe, 7.5 km ENE Ivohibe, 22.47°S , 46.96°E , 900 m, rainforest, 7.x.1997 (B.L.Fisher (Sylvain)); Fianarantsoa, Reserve Forestiere d'Agnalazaha , Mahabo, 42.9 km 215° Farafangana, 23.1938°S , 47.723°E , 20 m, littoral rainforest, 19.iv.2006 (B.L. Fisher et al.); Fianarantsoa, Soanierenana I Non Protected Area, 25.33 km SW Ambositra, 20.7214°S , 47.1099°E , 1723 m, savannah grassland, 6.ii.2010 (Andrianjaka Ravelomanana); Mahajanga, Reserve Speciale Marotandrano, Marotandrano 48.3 km S Mandritsara, 16.2832°S , 48.8144°E , 865 m, transition humid forest, 6.xii.2007 (B.L.Fisher et al.); Toamasina, 16 km S Moramanga, 19.0833°S , 48.2333°E , 950 m, rainforest, 18.xi.1990 (P. S. Ward); Toamasina, 19 km ESE Maroantsetra, 15.4833°S , 49.9°E , 350 m, rainforest, 22.iv.1989 (P. S. Ward); Toamasina, 5.3 km SSE Ambanizana, Andranobe, 15.6713°S , 49.974°E , 425 m, rainforest, 19.xi.1993 (B.L.Fisher); Toamasina, 6.3 km S Ambanizana, Andranobe, 15.6813°S , 49.958°E , 25 m, rainforest, 14.xi.1993 (B.L.Fisher); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroina, 15.5851°S , 50.0095°E , 825 m, rainforest, 2.xii.1993 (B.L.Fisher); Toamasina, Ambanizana, Parc National Masoala, 15.5717°S , 50.0061°E , 925 m, montane rainforest, 26.ii.2003 (D. Andriamalala, D. Silva, et al.); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.8496°S , 48.2947°E , 1010 m, montane rainforest, 3.iii.2007 (B.L.Fisher et al.); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.8394°S , 48.3084°E , 1080 m, montane rainforest, 4.iii.2007 (B.L.Fisher et al.); Toamasina, Analamay, 18.8062°S , 48.3371°E , 1068 m, montane rainforest, 21.iii.2004 (Malagasy ant team); Toamasina, Corridor Forestier Analamay-Mantadia, Ambatoharanana, 18.8042°S , 48.4008°E , 968 m, rainforest, 12.xii.2012 (B.L.Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Ambohibolakely, 18.779°S , 48.3638°E , 918 m, rainforest, 23.xi.2012 (B.L.Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Tsaravoniana, 18.7612°S , 48.4213°E , 939 m, rainforest, 2.xii.2012 (B.L.Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Tsaravoniana, 18.7646°S , 48.4194°E , 1039 m, rainforest, 2.xii.2012 (B.L.Fisher et al.); Toamasina, Foret Ambatovy, 14.3 km 57° Moramanga, 18.8508°S , 48.32°E , 1075 m, montane rainforest, 21.iii.2004 (Malagasy ant team); Toamasina, Ile Sainte Marie, Foret Kalalao, 9.9 km 34° Ambodifotatra, 16.9225°S , 49.8873°E , 100 m, rainforest, 24.xi.2005 (B.L.Fisher et al.); Toamasina, Montagne d'Akirindro 7.6 km 341° NNW Ambinanitelo, 15.2883°S , 49.5483°E , 600 m, rainforest, 17.iii.2003 (Fisher, Griswold et al.); Toamasina, Montagne d'Anjanaharibe , 18.0 km 21° NNE Ambinanitelo, 15.1883°S , 49.615°E , 470 m, rainforest, 8.iii.2003 (Fisher, Griswold et al.); Toamasina, Montagne d'Anjanaharibe , 19.5 km 27° NNE Ambinanitelo, 15.1783°S , 49.635°E , 1100 m, montane rainforest, 12.iii.2003 (Fisher, Griswold et al.); Toamasina, Parc National de Zahamena, Onibe River, 17.7591°S , 48.8547°E , 780 m, rainforest, 21.ii.2009 (B.L.Fisher et al.); Toamasina, Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, 17.743°S , 48.7294°E , 860 m, rainforest, 18.ii.2009 (B.L.Fisher et al.); Toamasina, Parc National Mananara-Nord, 7.1 km 261° Antanambe, 16.455°S , 49.7875°E , 225 m, rainforest, 14.xi.2005 (B.L.Fisher et al.); Toamasina, Parcelle K7 Tampolo, 17.2833°S , 49.4167°E , 10 m, littoral forest, 16.iv.2004 (Malagasy ant team); Toamasina, Parcelle K9 Tampolo, 17.175°S , 49.268°E , 10 m, littoral forest, 19.iv.2004 (Malagasy ant team); Toamasina, Res. Ambodiriana, 4.8 km 306°Manompana, along Manompana river, 16.6723°S , 49.7012°E , 125 m, rainforest, 18.xi.2005 (B.L.Fisher et al.); Toamasina, Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.8867°S , 49.2025°E , 520 m, rainforest, 1.xii.2005 (B.L.Fisher et al.); Toamasina, Reserve Speciale Ambatovaky, Sandrangato river, 16.8175°S , 49.295°E , 360 m, rainforest, 25.ii.2010 (B.L.Fisher et al.); Toamasina, S.F. Tampolo, 10 km NNE Fenoarivo Atn., 17.2825°S , 49.43°E , 10 m, littoral rainforest, 4.iv.1997 (B.L.Fisher); Toamasina, Sahafina forest 11.4 km W Brickaville, 18.8145°S , 48.962°E , 140 m, rainforest, 13.xii.2007 (B.L.Fisher et al.); Toamasina, Torotorofotsy, 18.8708°S , 48.3474°E , 1070 m, montane rainforest, marsh edge, 24.iii.2004 (Malagasy ant team); Toliara, 10 km NW Enakara, Res . Andohahela, 24.5667°S , 46.8167°E , 430 m, rainforest, 22.xi.1992 (B.L.Fisher); Toliara, 11 km NW Enakara, Res . Andohahela, 24.5667°S , 46.8333°E , 800 m, rainforest, 17.xi.1992 (B.L.Fisher); Toliara, 2.7 km WNW 302° Ste. Luce, 24.7717°S , 47.1717°E , 20 m, littoral rainforest, 9.xii.1998 (B.L.Fisher (J.-Baptiste)); Toliara, Anosy Region, Distric of Amboasary, 58Km SW of Fort Dauphin, 08 Km NW of Amboasary, Berenty Special Reserve, 25.0067°S , 46.3033°E , 85 m, Galery forest, 25.v.2003 ( Rin'ha , Mike); Toliara, Foret Classee d'Analavelona , 29.2 km 343° NNW Mahaboboka, 22.675°S , 44.19°E , 1100 m, montane rainforest, 18.ii.2003 (Fisher, Griswold et al.); Toliara, Foret Classee d'Analavelona , 29.4 km 343° NNW Mahaboboka, 22.675°S , 44.1867°E , 1050 m, montane rainforest, 21.ii.2003 (Fisher, Griswold et al.); Toliara, Foret Ivohibe 55.6 km N Tolagnaro, 24.5617°S , 47.2002°E , 650 m, rainforest, 4.xii.2006 (B.L.Fisher et al.); Toliara, Grand Lavasoa, 25.9 km W Tolagnaro, 25.0877°S , 46.749°E , 450 m, rainforest, 30.xi.2006 (B.L.Fisher et al.); Toliara, Mandena, 8.4 km NNE 30° Tolagnaro, 24.9517°S , 47.0017°E , 20 m, littoral rainforest, 20.xi.1998 (B.L.Fisher); Toliara, Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, 24.7585°S , 46.8537°E , 275 m, rainforest, 22.xi.2006 (B.L.Fisher et al.); Toliara, Res . Andohahela, 6 km SSW Eminiminy, 24.7333°S , 46.8°E , 330 m, rainforest, 4.ii.1993 (P. S. Ward); Toliara, Reserve Speciale d'Ambohijanahary , Foret d'Ankazotsihitafototra , 34.6 km 314° NW Ambaravaranala, 18.26°S , 45.4183°E , 1100 m, montane rainforest, 16.i.2003 (Fisher, Griswold et al.); Toliara, Reserve Speciale d'Ambohijanahary , Foret d'Ankazotsihitafototra , 35.2 km 312° NW Ambaravaranala, 18.2667°S , 45.4067°E , 1050 m, montane rainforest, 13.i.2003 (Fisher, Griswold et al.); Toliara, Reserve Speciale Kalambatritra, Ampanihy, 23.4635°S , 46.4631°E , 1270 m, montane rainforest, 9.ii.2009 (B.L.Fisher et al.); Bongolava Prefec. de Tsiroanomandidy, 6.xii.1974 (A.Peyrieras).