A revision of Malagasy species of Anochetus Mayr and Odontomachus Latreille (Hymenoptera: Formicidae). Author Fisher, B. L. Author Smith, M. A. text PLoS ONE 2008 3 1 23 http://dx.doi.org/10.1371/journal.pone.0001787 journal article 21401 Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species Forel Anochetus grandidieri Forel Figures: worker 3a-d, 5c; queen 3e-h; male 3i-j, 8b; map 6b Type material: Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species Forel, 1891: 108 Anochetus grandidieri Forel,1891: 108 [21]. Lectotype: worker, Madagascar Forest of the east coast M. Humblot Madagascar , Forest of the east coast ( M. Humblot ) ( MHNG ), present designation [examined], AntWeb CASENT0101819 . Brown, 1978: 606 [ 2 ] (description of worker) . Animalia Insecta Hymenoptera Formicidae Anochetus madecassus species Santschi, 1928: 54 Anochetus madecassus Santschi,1928: 54 [22]. Lectotype: dealate queen, Madagascar Descarpentries Lectotype Madagascar , Nossi-Be( Descarpentries ) ( NHMB ) Lectotype by present designation [examined] AntWeb CASENT0101098 . Synonymized with Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species grandidieri by Brown,1978: 557 [2]. Worker measurements: maximum and minimum based on all specimens, n = 20. HL 0.79-1.19, HW 0.71-1.06, CI 85-95, EL 0.08-0.13, ML 0.33-0.57, MI 41-54, SL 0.57-0.88, SI 78-86, WL 0.87-1.35, FL 0.57-0.90, PW 0.44-0.62. Figure 8. Animalia Insecta Hymenoptera Formicidae Anochetus genus Anochetus males, terminalia, lateral view. A, Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species boltoni CASENT0063847. B, Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species grandidieri CASENT0080660. B, Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species madagascarensis CASENT0063421. D, Animalia Insecta Hymenoptera Formicidae Anochetus pattersoni species pattersoni CASENT0172617. doi:10.1371/journal.pone.0001787.g008 Figure 9. NJ tree of K2P for five species of Animalia Insecta Hymenoptera Formicidae Anochetus genus Anochetus in Madagascar, Comoros and Aldabra (all specimens with.500 bp). Deep divergences evident between Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species madagascarensis , Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species grandidieri , and Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species goodmani are evident. Deep divergences within Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani are evident (In this tree, Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species A. boltoni falls within Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species goodmani ). The rightmost column of colors differentiate which biogeographical groupings of Wilme et al. [29] these populations fall. WCE-1 = Binara, Antsahabe. WCE-12 = Andavakoera, Ankarana. WCE-7= Kirindy Mite. WRDW-B = Vazimba, Androngonibe, Andranopasazy. doi:10.1371/journal.pone.0001787.g009 Table 2. Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species Anochetus goodmani within-species pair-wise partitioning of genetic variance for the CO1 DNA barcode. K2P distances are beneath the diagonal and the number substitutions are above the diagonal. Table 3. Comparison of the utility of various complimentary nuclear markers for species diagnosis in the ponerine ants of the Malagasy. Taxa 18S 28S ITS1 Comments Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species Anochetus goodmani Intra - no variation. nter - 2 bp from Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species A. boltoni , and 3 bp from Animalia Insecta Hymenoptera Formicidae Odontomachus troglodytes species O. troglodytes , Animalia Insecta Hymenoptera Formicidae Odontomachus coquereli species O. coquereli Intra - no variation across north nter -.15 bp divergent from Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species A. madagascarensis . Intra - extreme variation (length and substitution) across range. Some corresponding to deep CO1 splits - provisionally orthologous. However, deep paralogous divergences have been sequenced within single individuals through different amplifications and extractions. rRNA is, a priori, difficult to differentiate orthologous from paraologous. Not as immediately useful as an independent marker without cloning. Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species Anochetus boltoni Intra - no variation. nter - 2 bp from Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani , 2 bp from Animalia Insecta Hymenoptera Formicidae Odontomachus troglodytes species O. troglodytes and no difference from Animalia Insecta Hymenoptera Formicidae Odontomachus coquereli species O. coquereli . N/A Intraspecific variation of 1% (indels and substitutions) between the two sampled populations. Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species Anochetus madagascarensis N/A Intra - no variation. nter -.15 bp divergent from Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani . Intra - variation that does NOT reflect CO1 variation. rRNA is, a priori, difficult to differentiate orthologous from paraologous. Not as immediately useful as an independent marker without cloning. -Positive Wolbachia test. Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species Anochetus grandidieri N/A N/A Low iIntraspecific variation that does reflect CO1 geographic variation. - Positive Wolbachia test. Animalia Insecta Hymenoptera Formicidae Anochetus pattersoni species Anochetus pattersoni N/A N/A N/A Animalia Insecta Hymenoptera Formicidae Odontomachus coquereli species Odontomachus coquereli Intra - no variation. nter - 2 bp from Animalia Insecta Hymenoptera Formicidae Odontomachus troglodytes species O. troglodytes . 2bp from Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species A. boltoni ,and 3 bp from Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani . Intra - variation. Large variation at geographically distal ends of distribution. nter - differentiates between three Malagasy species. Intraspecific variation that only partially reflects geography and CO1 variation - while some clearly does not. Paralogous and orthologous Animalia Insecta Hymenoptera Formicidae Odontomachus troglodytes species Odontomachus troglodytes Intra - no variation. nter - 2 bp from Animalia Insecta Hymenoptera Formicidae Odontomachus coquereli species O. coquereli . 3 bp from Animalia Insecta Hymenoptera Formicidae Anochetus boltoni species A. boltoni ,and3bp from Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani . Intra - some variation that does not correspond to geography or CO1. nter - does not differentiate between Animalia Insecta Hymenoptera Formicidae Odontomachus simillimus species O. simillimus Intraspecific variation that only partially reflects geography and CO1 variation All specimens tested positive for Wolbachia. Animalia Insecta Hymenoptera Formicidae Odontomachus simillimus species Odontomachus simillimus N/A Intra - no variation. nter - does not differentiate Animalia Insecta Hymenoptera Formicidae Odontomachus troglodytes species O. troglodytes N/A doi:10.1371/journal.pone.0001787.t003 Queen measurements: maximum and minimum based on n = 5. HL 0.88-1.15, HW 0.81-1.07, CI 92-96, EL 0.17-0.23, ML 0.39-0.56, MI 44-49, SL 0.62-0.87, SI 77-81, WL 1.08- 1.46. FL 0.68-0.96, PW 0.60-0.78. Male measurements: maximum and minimum based on n = 5 from Madagascar: HL 0.58-0.73, HW 0.78-0.94, CI 129- 135, EL 0.37-0.46, SL 0.10-0.15, SI 13-16, WL 1.17-1.52, FL 0.78-1.08 Worker diagnosis: Inner blade of mandible without teeth and denticles; apical end of inner blade without a notched semicircular concavity (Fig. 2a). Eyes small (0.05-0.11 mm), projecting dorsolaterally. In full face view, antennal scape usually not reaching, and not surpassing posterior margin of occipital lobe. Dorsal surface of head with numerous short setae. Pilosity and sculpture as in Figures 3 a-d. Queens alate: Very similar to workers, only slightly larger than respective size class (Figs 3e-h). Ergatoid queens not recorded. Within a single locality, two size classes of workers, queens and males are present in this species, but the differences within a site do not hold up when variation across all sites is included. These differences suggest that two reproductive and developmental pathways can occur in this species. Further work is needed to explore the biotic or abiotic factors that trigger the development of small and large castes. The species is most similar to Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species A. madagascarensis but can be easily distinguished by its small eyes and scape that does not surpass the occipital lobe. Animalia Insecta Hymenoptera Formicidae Anochetus madagascarensis species A. madagascarensis has large eyes (0.24-0.26 mm), and scapes that surpass occipital lobes. Distribution and biology. Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A. grandidieri is endemic to Madagascar and is widespread throughout Madagascar in forest and shrubland habitats below 1,550 m elevation (Fig 4b). It has been collected in gallery, dry, littoral, lowland, and montane forest, in desert spiny bush thicket in the southwest, and Uapaca woodland in the ceIntral plateau. As in many soil dwelling ants, Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A. grandidieri has reduced eyes (EL/HW 0.11-0.13) and short scapes. Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A. grandidieri is the only Animalia Insecta Hymenoptera Formicidae Anochetus genus Anochetus in Madagascar with these soil nesting modifications. The subterranean habitat of this species may allow it to survive in a wide range of habitats in Madagascar from desert to woodland to montane forest. Out of 453 collecting events, Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A. grandidieri was most often recorded in sifted litter (97 collection records), rotten logs (96), and Malaise traps (155). CO1. Shallow iIntraspecific (average within species sequence divergence of 2.72, SE = 0.048) and deep interspecific divergences(9.4% SE = 0.05) between Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A. grandidieri and the other species. Small and large castes had identical DNA barcodes. (Figs 9, 16). Diagnostic barcoding loci. Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species A grandidieri : T-273, T-282, T- 306, A-312, (shared with one population of Animalia Insecta Hymenoptera Formicidae Anochetus goodmani species A. goodmani ), A-312, T- 333, A-483, T-528 (all 3rd base pair positions). Specimens examined for Animalia Insecta Hymenoptera Formicidae Anochetus grandidieri species Anochetus grandidieri : Specimens from 456 separate collection events from the following 140 localities were examined. MADAGASCAR Province Antsiranana Sakalava Beach MADAGASCAR : Province Antsiranana : Sakalava Beach ; MADAGASCAR Montagne des Francais Antsiranana Montagne des Francais , 7.2 km 142° SE Antsiranana ; 49.1725 -12.5283 Madagascar Antsiranana II Pref: Antsahampano S.-Pref Montagne 2004 11 16 D.Lees, R.Ranaivosolo & P.Razafindraibe Antsiranana II Pref: Antsahampano S.-Pref : Montagne d'Ambre. Site MD2 ; 49.1795 -12.53444 Madagascar Parc National Montagne d'Ambre Joffreville 2001 01 20 Fisher, Griswold et al. Parc National Montagne d'Ambre , 3.6 km 235° SW Joffreville ; 49.22583 -12.86361 Madagascar Reserve Speciale de l'Ankarana Anivorano Nord 2001 02 16 Fisher, Griswold et al. Reserve Speciale de l'Ankarana , 13.6 km 192° SSW Anivorano Nord ; 49.7 -12.97 Madagascar Foret d'Ampondrabe Daraina 2003 12 11 B.L.Fisher Foret d'Ampondrabe , 26.3 km 10° NNE Daraina ; 49.55667 -13.21167 Madagascar Foret d' Antsahabe Daraina 2003 12 12 B.L.Fisher Foret d' Antsahabe , 11.4 km 275° W Daraina ; 49.61667 -13.255 Madagascar Foret de Binara Daraina 2003 12 01 B.L.Fisher Foret de Binara , 7.5 km 230° SW Daraina ; 49.60333 -13.26333 Madagascar Foret de Binara Daraina 2004 11 19 B.L.Fisher Foret de Binara , 9.1 km 233° SW Daraina ; 48.30722 -13.41639 Madagascar Foret de Binara Nosy Be 1992 08 13 Gary D. Alpert Nosy Be , Lokobe Forest ; 48.55167 -13.4645 Madagascar Foret Ambato Ambanja 2004 12 08 B.L.Fisher Foret Ambato , 26.6 km 33° Ambanja ; 50.10167 -13.71533 Madagascar Ambondrobe Vohemar 2004 11 29 B.L.Fisher Ambondrobe , 41.1 km 175° Vohemar ; 48.16167 -13.79861 Madagascar Ampasindava, Foret d'Ambilanivy Ambaliha 2001 03 04 Fisher, Griswold et al. Ampasindava, Foret d'Ambilanivy , 3.9 km 181° SAmbaliha ; 48.43333 -13.96167 Madagascar R.S. Manongarivo Antanambao 1998 11 08 B.L.Fisher R.S. Manongarivo , 10.8 km 229° SW Antanambao ; 48.42333 -13.97667 Madagascar R.S. Manongarivo Antanambao 1998 10 11 B.L.Fisher R.S. Manongarivo , 12.8 km 228° SW Antanambao ; 48.42833 -13.99833 Madagascar R.S. Manongarivo Antanambao 1998 10 19 B.L.Fisher R.S. Manongarivo , 14.5 km 220° SW Antanambao ; 47.91433 -14.30889 Madagascar Foret d'Anabohazo Maromandia 2001 03 11 Fisher, Griswold et al. Foret d'Anabohazo , 21.6 km 247° WSW Maromandia ; 49.76 -14.435 Madagascar Parc National de Marojejy Andapa 2003 11 15 B.L.Fisher et al. Parc National de Marojejy , Manantenina River , 27.6 km 35° NE Andapa , 9.6 km 327° NNW Manantenina ; 49.775 -14.43667 Madagascar Parc National de Marojejy, Manantenina River Andapa 2003 11 12 B.L.Fisher et al. Parc National de Marojejy, Manantenina River , 28.0 km 38° NE Andapa , 8.2 km 333° NNW Manantenina ; 49.774 -14.43817 Madagascar Parc National Marojejy Parc National Marojejy 2005 05 30 Rin'Ha, Mike Parc National Marojejy ; 49.78564 -14.44533 Madagascar Marojejy Marojejy 1993 11 23 Gary D. Alpert Marojejy R.N.I. #12 ; 50.18367 -14.67933 Madagascar Foret Ambanitaza Antalaha 2004 11 26 B.L.Fisher Foret Ambanitaza , 26.1km 347° Antalaha ; 49.46667 -14.75 Madagascar Res. Anjanaharibe-Sud Befingotra 1994 11 09 B.L.Fisher 9.2 km WSW Befingotra , Res. Anjanaharibe-Sud ; 49.5 -14.75 Madagascar Res. Anjanaharibe-Sud Befingotra 1994 10 19 B.L.Fisher 6.5 km SSW Befingotra , Res. Anjanaharibe-Sud ; 49.51019 -14.75775 Madagascar Res. d' Anjanaharibe-Sud Andapa 1994 11 02 Gary D. Alpert 17 km W Andapa , Res. d' Anjanaharibe-Sud ; 50.26167 -14.93806 Madagascar 9.2 km WSW Befingotra Antalaha 1993 12 06 Alpert et. al. 5 km SW Antalaha ; 50.33803 -15.29128 Madagascar 14 km W Cap Est Ambato 1994 11 12 Gary D. Alpert 14 km W Cap Est , Ambato ; 50.27028 -15.69694 Madagascar Fotodriana Cap Masoala 1993 11 28 Gary D. Alpert Fotodriana , Cap Masoala . 45.90833 -16.05167 Madagascar Province Mahajanga Mahavavy River 2002 12 01 Fisher, Griswold et al. Province Mahajanga : Mahavavy River , 6.2 km 145° SE Mitsinjo ; 47.04861 -16.26722 Madagascar Reserve d'Ankoririka Tsaramandroso 2001 04 09 Fisher, Griswold et al. Reserve d'Ankoririka , 10.6 km 13° NE de Tsaramandroso ; 46.81333 -16.31944 Madagascar Ampijoroa National Park Maevatanana 2003 11 09 Rin'ha Harin'Hala Ampijoroa National Park , 160 km N Maevatanana , Mahajanga Prov ., deciduous forest ; 45.32667 -16.37667 Madagascar Parc National de Namoroka Vilanandro 2002 11 08 Fisher, Griswold et al. Parc National de Namoroka , 17.8 km 329° WNW Vilanandro ; 45.31 -16.40667 Madagascar Parc National de Namoroka Vilanandro 2002 11 12 Fisher, Griswold et al. Parc National de Namoroka , 16.9 km 317° NW Vilanandro ; 45.35 -16.46667 Madagascar Parc National de Namoroka Vilanandro 2002 11 04 Fisher, Griswold et al. Parc National de Namoroka , 9.8 km 300° WNW Vilanandro ; 44.36833 -16.925 Madagascar Reserve Speciale de Bemarivo Besalampy 2002 11 19 Fisher, Griswold et al. Reserve Speciale de Bemarivo , 23.8 km 223° SW Besalampy ; 44.71817 -18.70944 Madagascar Parc National Tsingy de Bemaraha Antsalova 2001 11 16 Fisher-Griswold Arthropod Team Parc National Tsingy de Bemaraha , 10.6 km ESE 123° Antsalova ; 44.44067 -19.02139 Madagascar Foret de Tsimembo Soatana 2001 11 21 Fisher-Griswold Arthropod Team Foret de Tsimembo , 8.7 km 336° NNW Soatana ; 44.81467 -19.13222 Madagascar Parc National Tsingy de Bemaraha Bekopaka 2001 11 11 Fisher-Griswold Arthropod Team Parc National Tsingy de Bemaraha , 2.5 km 62° ENE Bekopaka , Ankidrodroa River ; 44.828 -19.14194 Madagascar Parc National Tsingy de Bemaraha Bekopaka 2001 11 06 Fisher-Griswold Arthropod Team Parc National Tsingy de Bemaraha , 3.4 km 93° E Bekopaka , Tombeau Vazimba ; 49.635 -15.17833 Madagascar Province Toamasina Ambinanitelo 2003 03 12 Fisher, Griswold et al. Province Toamasina : Montagne d'Anjanaharibe , 19.5 km 27° NNE Ambinanitelo ; 49.615 -15.18833 Madagascar Montagne d'Anjanaharibe Ambinanitelo 2003 03 08 Fisher, Griswold et al. Montagne d'Anjanaharibe , 18.0 km 21 ° NNE Ambinanitelo ; 49.54833 -15.28833 Madagascar Montagne d'Akirindro Ambinanitelo 2003 03 17 Fisher, Griswold et al. Montagne d'Akirindro 7.6 km 341 ° NNW Ambinanitelo ; 49.9 -15.48333 Madagascar 19 km ESE Maroantsetra Maroantsetra 1989 04 22 P. S. Ward 19 km ESE Maroantsetra ; 50 -15.56667 Madagascar 6.9 km NE Ambanizana Ambohitsitondroina 1993 12 02 B.L.Fisher 6.9 km NE Ambanizana , Ambohitsitondroina ; 50.00611 -15.57167 Madagascar Ambanizana Parc National Masoala 2003 02 26 D. Andriamalala, D. Silva, et al. Ambanizana , Parc National Masoala ; 49.96667 -15.66667 Madagascar 5.3 km SSE Ambanizana Andranobe 1993 11 21 B.L.Fisher 5.3 km SSE Ambanizana , Andranobe ; 49.95 -15.68333 Madagascar 5.3 km SSE Ambanizana Andranobe 1993 11 14 B.L.Fisher 6.3 km S Ambanizana , Andranobe ; 1 km W Andampibe , Cap Masoala ; 49.7875 -16.455 Madagascar Parc National Mananara-Nord Antanambe 2005 11 15 B.L.Fisher et al. Parc National Mananara-Nord , 7.1 km 261° Antanambe ; MADAGASCAR Foret d'Analava Mandrisy Antanambe Foret d'Analava Mandrisy , 5.9 km 195° Antanambe ; MADAGASCAR Res. Ambodiriana Manompana Res. Ambodiriana , 4.8 km 306° Manompana , along Manompana River ; 49.96417 -16.82433 Madagascar Ile Sainte Marie Foret Ambohidena 2005 11 21 B.L.Fisher et al. Ile Sainte Marie , Foret Ambohidena , 22.8 km 44° Ambodifotatra ; 49.88733 -16.9225 Madagascar Ile Sainte Marie Foret Kalalao 2005 11 24 B.L.Fisher et al. Ile Sainte Marie , Foret Kalalao , 9.9 km 34° Ambodifotatra ; 49.43333 -17.28333 Madagascar Parcelle E3 Tampolo Tampolo 2004 04 14 Malagasy ant team Parcelle E3 Tampolo ; 49.41667 -17.28333 Madagascar S.F. Tampolo, Fenoarivo Atn 1997 04 03 B.L.Fisher S.F. Tampolo, 10 km NNE Fenoarivo Atn . ; 49.46667 -17.65 Madagascar Bridge at Onibi Mahavelona 1993 12 01 A. Pauly Bridge at Onibi , NW of Mahavelona ; 49.5 -17.66667 Madagascar Mahavelona Foulpointe 1995 01 01 A. Pauly Mahavelona ( Foulpointe ) ; 49.49583 -17.67017 Madagascar Mahavelona Mahavelona 2005 11 27 B.L.Fisher et al. 2.1 km 315° Mahavelona ; 49.51667 -17.68333 Madagascar Foulpointe Foulpointe 1993 11 05 A.Pauly Foulpointe ; 49.2025 -17.88667 Madagascar Reserve Betampona Camp Vohitsivalana 2005 12 03 B.L.Fisher et al. Reserve Betampona , Camp Vohitsivalana , 37.1 km 338° Toamasina ; 49.19967 -17.924 Madagascar Reserve Betampona Camp Rendrirendry 2005 11 29 B.L.Fisher et al. Reserve Betampona , Camp Rendrirendry 34.1 km 332° Toamasina ; 49.09167 -18.04833 Madagascar F.C. Sandranantitra F.C. Sandranantitra 1999 01 18 H.J.Ratsirarson F.C. Sandranantitra ; 48.57833 -18.19833 Madagascar F.C. Didy F.C. Didy 1998 12 16 H.J.Ratsirarson F.C. Didy ; 48.81333 -18.695 Madagascar F.C. Andriantantely F.C. Andriantantely 1998 12 04 H.J.Ratsirarson F.C. Andriantantely ; 48.42667 -18.79167 Madagascar P.N. Mantadia P.N. Mantadia 1998 11 25 H.J.Ratsirarson P.N. Mantadia ; 48.33707 -18.80623 Madagascar Analamay Analamay 2004 03 23 Malagasy ant team Analamay ; 48.32 -18.85083 Madagascar Foret Ambatovy Moramanga 2004 12 18 B.L.Fisher Foret Ambatovy , 14.3 km 57° Moramanga ; 48.34737 -18.87082 Madagascar Torotorofotsy Torotorofotsy 2004 03 29 Malagasy ant team Torotorofotsy ; MADAGASCAR Andasibe National Park West of ANGAP office Andasibe National Park , botanic garden near eIntrance, West of ANGAP office ; 48.41833 -18.92778 Madagascar Res. Analamazaotra Andasibe 2006 01 07 A.Ballerio Res. Analamazaotra , Parc National , Andasibe ; 46.56333 -20.59333 Madagascar Fianarantsoa Foret 2003 01 22 Fisher, Griswold et al. Fianarantsoa : Foret d'Atsirakambiaty, 7.6 km 285° WNW Itremo ; 47 -21 Madagascar Ranomafana Nat. Park Miaranony Forest 1992 10 26 Emile Rajeriarison Ranomafana Nat. Park , Miaranony Forest ; 47.36983 -21.22617 Madagascar Vohiparara broken bridge Fianarantsoa Prov 2002 02 26 R. Harin'Hala Vohiparara broken bridge , Fianarantsoa Prov . ; 47.39667 -21.23667 Madagascar Parc National de Ranomafana Sahamalaotra River 2003 03 31 Fisher, Griswold et al. Parc National de Ranomafana , Sahamalaotra River , 6.6 km 310° NW Ranomafana ; 47.42667 -21.24833 Madagascar Parc Nationale Ranomafana Talatakely 1998 04 09 CE Griswold, DH Kavanaugh, ND Penny, MJ Raherilalao, JS Ranorianarisoa, J Schwei Parc Nationale Ranomafana : Talatakely ; 47.41667 -21.25 Madagascar Ranomafana Ifandiana 1989 04 27 P.S.Ward 3 km W Ranomafana , nr. Ifandiana ; 47.41945 -21.25041 Madagascar Ranomafana National Park Talatakely 1998 04 11 ME Irwin & EI Schlinger research cabin at Talatakely , Ranomafana National Park ; 47.40717 -21.25083 Madagascar Ranomafana National Park Fianarantsoa Prov 2003 05 17 R. Harin'Hala radio tower, Ranomafana National Park , Fianarantsoa Prov . ; 47.42178 -21.25833 Madagascar Ranomafana National Park Namorona River 1998 04 11 ME Irwin & EI Schlinger Namorona River at footbridge, Ranomafana National Park ; 47.425 -21.26583 Madagascar Ranomafana National Park Ranomafana National Park, Tavolo tree 1996 04 08 Alpert et. al. Ranomafana National Park , Tavolo tree ; 47.42017 -21.2665 Madagascar Belle Vue trail Ranomafana National Park 2003 03 10 R. Harin'Hala Belle Vue trail , Ranomafana National Park , Fianarantsoa Prov . ; 47.41667 -21.26667 Madagascar Belle Vue trail Ranomafana 1993 09 23 M.Stebbine, W.E.Steiner, et. al. 7 km W Ranomafana ; 47.4304 -21.28955 Madagascar Vatoharanana Vatoharanana 2003 03 30 V.C.Clark Vatoharanana ; 47.43333 -21.29 Madagascar Parc National de Vatoharanana River 2003 03 27 Fisher, Griswold et al. Parc National deRanomafana , Vatoharanana River , 4.1 km 231 ° SW Ranomafana ; 47.42615 -21.29073 Madagascar P.N. Ranomafana Vatoharanana-Ankovoka 2003 03 13 V. Clark P.N. Ranomafana , Vatoharanana-Ankovoka ; 47.94194 -21.39306 Madagascar 8 km E Kianjavato Vatovavy Forest 1993 12 11 Alpert 8 km E Kianjavato , Vatovavy Forest ; MADAGASCAR 8 km E Kianjavato Kianjavato 7.6 km 122° Kianjavato , Foret Classee Vatovavy ; 47.41 -21.51167 Madagascar 8 km E Kianjavato Andrambovato 2005 06 03 B.L.Fisher et al. 2 km W Andrambovato , along river Tatamaly ; 47.33889 -21.8275 Madagascar Foret d'Ambalagoavy Nord Ikongo 2000 12 01 R. Harin'Hala & M.E. Irwin Foret d'Ambalagoavy Nord , Ikongo , Ambatombe ; 47.01667 -22.21667 Madagascar 45 km S. Ambalavao Ambalavao 1993 09 25 B.L.Fisher 45 km S. Ambalavao ; 47.01667 -22.21667 Madagascar 45 km S Ambalavao Ambalavao 1993 10 31 B.L.Fisher 45 km S Ambalavao ; 47 -22.23333 Madagascar Res. Andringitra Ambalavao 1993 10 10 B.L.Fisher 43 km S Ambalavao , Res. Andringitra ; 45.35167 -22.29833 Madagascar Parc National d'Isalo Ambovo Springs 2003 02 09 Fisher, Griswold et al. Parc National d'Isalo , Ambovo Springs , 29.3 km 4° N Ranohira ; 46.89833 -22.42167 Madagascar 8.0 km NE Ivohibe 8.0 km NE Ivohibe 1997 11 03 B.L.Fisher (Sylvain) 8.0 km NE Ivohibe ; 46.93833 -22.42667 Madagascar 9.0 km NE Ivohibe 9.0 km NE Ivohibe 1997 11 12 B.L.Fisher (Sylvain) 9.0 km NE Ivohibe ; 46.96 -22.47 Madagascar R.S. Ivohibe Ivohibe 1997 10 07 B.L.Fisher (Sylvain) R.S. Ivohibe , 7.5 km ENE Ivohibe ; 45.46167 -22.48167 Madagascar Parc National d'Isalo Ranohira 2003 01 27 Fisher, Griswold et al. Parc National d'Isalo , 9.1 km 354° N Ranohira ; 45.12833 -22.59167 Madagascar Foret d'Analalava Ranohira 2003 02 01 Fisher, Griswold et al. Foret d'Analalava , 29.6 km 280° W Ranohira ; 47.18183 -22.791 Madagascar Foret de Vevembe Farafangana 2006 04 23 B.L. Fisher et al. Foret de Vevembe , 66.6 km 293° Farafangana ; MADAGASCAR Province Toliara Province Toliara : 45.41833 -18.26 Madagascar Reserve Speciale d'Ambohijanahary Foret d'Ankazotsihitafototra 2003 01 16 Fisher, Griswold et al. Reserve Speciale d'Ambohijanahary , Foret d'Ankazotsihitafototra , 34.6 km 314° NW Ambaravaranala ; 45.40667 -18.26667 Madagascar Reserve Speciale d'Ambohijanahary Foret d'Ankazotsihitafototra 2003 01 13 Fisher, Griswold et al. Reserve Speciale d'Ambohijanahary , Foret d'Ankazotsihitafototra , 35.2 km 312° NW Ambaravaranala ; 44.85 -22.46667 Madagascar Vohibasia Forest Sakaraha 1996 01 13 B.L.Fisher Vohibasia Forest , 59 km NE Sakaraha ; 44.85 -22.46667 Madagascar southern Isoky-Vohimena Forest Sakaraha 1996 01 21 B.L.Fisher southern Isoky-Vohimena Forest , 59 km NE Sakaraha ; 44.17167 -22.64333 Madagascar Foret Classee d'Analavelona Mahaboboka 2003 02 22 Fisher, Griswold et al. Foret Classee d'Analavelona , 33.2 km 344° NNW Mahaboboka ; 44.19 -22.675 Madagascar Foret Classee d'Analavelona Mahaboboka 2003 02 18 Fisher, Griswold et al. Foret Classee d'Analavelona , 29.2 km 343° NNW Mahaboboka ; 44.18667 -22.675 Madagascar Foret Classee d'Analavelona Mahaboboka 2003 02 21 Fisher, Griswold et al. Foret Classee d'Analavelona , 29.4 km 343° NNW Mahaboboka ; 43.42067 -22.80222 Madagascar Foret de Tsinjoriaky Tsifota 2002 03 06 Fisher-Griswold Arthropod Team Foret de Tsinjoriaky , 6.2 km 84° E Tsifota ; 44.71 -22.84333 Madagascar Sakaraha Parc National de Zombitse 2003 02 05 Fisher, Griswold et al. Parc National de Zombitse , 19.8 km 84° E Sakaraha ; 44.70167 -22.88833 Madagascar Parc National de Zombitse Sakaraha 2003 02 08 Fisher, Griswold et al. Parc National de Zombitse , 17.7 km 98° E Sakaraha ; 44.68333 -22.9 Madagascar 15 km E Sakaraha Sakaraha 1993 02 15 P.S.Ward 15 km E Sakaraha ; 44.12133 -23.52417 Madagascar Foret de Mite Tongobory 2002 02 27 Fisher-Griswold Arthropod Team Foret de Mite , 20.7 km 29° WNW Tongobory ; 44.63333 -23.65 Madagascar Sept Lacs; Beza-Mahafaly, 27 km E Betioky Betioky 1997 04 23 B.L.Fisher Sept Lacs; Beza-Mahafaly, 27 km E Betioky ; 44.63333 -23.68333 Madagascar Ehazoara Canyon Betioky 1997 04 27 B.L.Fisher Ehazoara Canyon , 26 km E Betioky ; 47.21944 -24.43806 Madagascar Tolanaro Mahermano Mt 1995 01 25 K.C.Emberton 70.7 km NNE Tolanaro , Mahermano Mt .; 11 km NW Enakara , Res. Andohahela ; 46.81667 -24.56667 Madagascar 10 km NW Enakara Res. Andohahela 1992 11 22 B.L.Fisher 10 km NW Enakara , Res. Andohahela ; 46.8 -24.73333 Madagascar Andohahela Res 1993 02 04 P. S. Ward Res . Andohahela , 6 km SSW Eminiminy ; 46.75167 -24.76389 Madagascar Parc National d'Andohahela Col du Sedro 2002 01 21 Fisher-Griswold Arthropod Team Parc National d'Andohahela , Col du Sedro , 3.8 km 113° ESE Mahamavo , 37.6 km 341° NNW Tolagnaro ; 46.76683 -24.76389 Madagascar Parc National d'Andohahela Manampanihy River 2002 01 24 Fisher-Griswold Arthropod Team Parc National d'Andohahela , Manampanihy River , 5.4 km 113° ESE Mahamavo , 36.7 km 343° NNW Tolagnaro ; 47.17167 -24.77167 Madagascar 2.7 km WNW 302° Ste. Luce Ste. Luce 1998 12 09 B.L.Fisher (J.-Baptiste) 2.7 km WNW 302° Ste. Luce ; MADAGASCAR 9.2 km N Tolanaro Ilapany Mt 9.2 km N Tolanaro , Ilapany Mt . ; 46.75361 -24.93694 Madagascar 9.2 km N Tolanaro Tolanaro 1995 02 07 K.C.Emberton et al 29.5 km WNW Tolanaro , Vasiha Mt . ; 46.6455 -24.93 Madagascar Parc National d'Andohahela Foret d'Ambohibory 2002 01 16 Fisher-Griswold Arthropod Team Parc National d'Andohahela , Foret d'Ambohibory , 1.7 km 61° ENE Tsimelahy , 36.1 km 308° NW Tolagnaro ; 47.00167 -24.95167 Madagascar Mandena Tolagnaro 1998 11 20 B.L.Fisher Mandena , 8.4 km NNE 30° Tolagnaro ; 46.2715 -24.95694 Madagascar Reserve Prive Berenty Foret de Bealoka 2002 02 03 Fisher-Griswold Arthropod Team Reserve Prive Berenty , Foret de Bealoka , Mandrare River , 14.6 km 329° NNW Amboasary ; 46.306 -25.00778 Madagascar Reserve Prive Berenty Foret de Malaza 2002 02 06 Fisher-Griswold Arthropod Team Reserve Prive Berenty , Foret de Malaza , Mandrare River , 8.6 km 314° NW Amboasary ; 46.3 -25.01667 Madagascar Reserve Berenty Réserve Berenty 1993 02 09 P. S. Ward Reserve Berenty; Foret de Petriky , 12.5 km W 272° Tolagnaro ; 4.4 km 148° SSE Lavanono ; 45.14683 -25.59444 Madagascar Reserve Speciale de Cap Sainte Marie Marovato 2002 02 13 Fisher-Griswold Arthropod Team Reserve Speciale de Cap Sainte Marie , 14.9 km 261° W Marovato ; 44.73117 -22.8405 Madagascar Reserve Speciale de Cap Sainte Marie Zombitse National Park 2001 12 16 R. Harin'Hala near road, Zombitse National Park , Tulear Prov .; near ANGAP office, ZombitseNational Park , Tulear Prov .; 44.591 -23.6865 Madagascar Parcel I, Beza Mahafaly Reserve Tulear Province 2001 10 15 R. Harin'Hala Parcel I, Beza Mahafaly Reserve , near research station, Tulear Province ; 46.62667 -24.93683 Madagascar Tsimelahy - Parcel II Andohahela National Park 2002 11 26 M.E. Irwin, F.D. Parker, R. Harin'Hala Tsimelahy - Parcel II , Andohahela National Park , transition forest, Tulear Province .