Anochetus grandidieri Forel HNS
Figures: worker 3a-d, 5c; queen 3e-h; male 3i-j, 8b; map 6b
Type material:
Anochetus grandidieri Forel, 1891: 108 HNS [21]. Lectotype: worker, Madagascar, Forest of the east coast (M. Humblot) (MHNG), present designation [examined], AntWeb CASENT0101819. Brown, 1978: 606 [2] (description of worker) .
Anochetus madecassus Santschi, 1928: 54 HNS [22]. Lectotype: dealate queen, Madagascar, Nossi-Be( Descarpentries) (NHMB) Lectotype by present designation [examined] AntWeb CASENT0101098. Synonymized with grandidieri HNS 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.
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 A. madagascarensis HNS but can be easily distinguished by its small eyes and scape that does not surpass the occipital lobe. A. madagascarensis HNS has large eyes (0.24-0.26 mm), and scapes that surpass occipital lobes.
Distribution and biology. A. grandidieri HNS 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, A. grandidieri HNS has reduced eyes (EL/HW 0.11-0.13) and short scapes . A. grandidieri HNS is the only Anochetus HNS 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, A. grandidieri HNS 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 A. grandidieri HNS and the other species. Small and large castes had identical DNA barcodes. (Figs 9, 16).
Diagnostic barcoding loci. A grandidieri HNS: T-273, T-282, T- 306, A-312, (shared with one population of A. goodmani HNS), A-312, T- 333, A-483, T-528 (all 3rd base pair positions).
Specimens examined for Anochetus grandidieri HNS: Specimens from 456 separate collection events from the following 140 localities were examined. MADAGASCAR: Province Antsiranana: Sakalava Beach; Montagne des Francais, 7.2 km 142° SE Antsiranana; Antsiranana II Pref: Antsahampano S.-Pref: Montagne d'Ambre. Site MD2 ; Parc National Montagne d'Ambre, 3.6 km 235° SW Joffreville; Reserve Speciale de l'Ankarana, 13.6 km 192° SSW Anivorano Nord; Foret d'Ampondrabe, 26.3 km 10° NNE Daraina; Foret d' Antsahabe, 11.4 km 275° W Daraina; Foret de Binara, 7.5 km 230° SW Daraina; Foret de Binara, 9.1 km 233° SW Daraina; Nosy Be, Lokobe Forest; Foret Ambato, 26.6 km 33° Ambanja; Ambondrobe, 41.1 km 175° Vohemar; Ampasindava, Foret d'Ambilanivy, 3.9 km 181° S Ambaliha; R.S. Manongarivo, 10.8 km 229° SW Antanambao; R.S. Manongarivo, 12.8 km 228° SW Antanambao; R.S. Manongarivo, 14.5 km 220° SW Antanambao; Foret d'Anabohazo, 21.6 km 247° WSW Maromandia; Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina; Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina; Parc National Marojejy; Marojejy R.N.I. #12 ; Foret Ambanitaza, 26.1km 347° Antalaha; 9.2 km WSW Befingotra, Res. Anjanaharibe-Sud ; 6.5 km SSW Befingotra, Res. Anjanaharibe-Sud ; 17 km W Andapa, Res. d' Anjanaharibe-Sud ; 5 km SW Antalaha; 14 km W Cap Est, Ambato; Fotodriana, Cap Masoala. Province Mahajanga: Mahavavy River, 6.2 km 145° SE Mitsinjo; Reserve d'Ankoririka, 10.6 km 13° NE de Tsaramandroso; Ampijoroa National Park, 160 km N Maevatanana, Mahajanga Prov., deciduous forest ; Parc National de Namoroka, 17.8 km 329° WNW Vilanandro; Parc National de Namoroka, 16.9 km 317° NW Vilanandro; Parc National de Namoroka, 9.8 km 300° WNW Vilanandro; Reserve Speciale de Bemarivo, 23.8 km 223° SW Besalampy; Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova; Foret de Tsimembo, 8.7 km 336° NNW Soatana; Parc National Tsingy de Bemaraha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River; Parc National Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba; Province Toamasina: Montagne d'Anjanaharibe, 19.5 km 27° NNE Ambinanitelo; Montagne d'Anjanaharibe, 18.0 km 21 ° NNE Ambinanitelo; Montagne d'Akirindro 7.6 km 341 ° NNW Ambinanitelo; 19 km ESE Maroantsetra; 6.9 km NE Ambanizana, Ambohitsitondroina; Ambanizana, Parc National Masoala; 5.3 km SSE Ambanizana, Andranobe; 6.3 km S Ambanizana, Andranobe; 1 km W Andampibe, Cap Masoala; Parc National Mananara-Nord, 7.1 km 261° Antanambe; Foret d'Analava Mandrisy, 5.9 km 195° Antanambe; Res. Ambodiriana, 4.8 km 306° Manompana, along Manompana River; Ile Sainte Marie, Foret Ambohidena, 22.8 km 44° Ambodifotatra; Ile Sainte Marie, Foret Kalalao, 9.9 km 34° Ambodifotatra; Parcelle E3 Tampolo; S.F. Tampolo, 10 km NNE Fenoarivo Atn. ; Bridge at Onibi, NW of Mahavelona; Mahavelona (Foulpointe) ; 2.1 km 315° Mahavelona; Foulpointe; Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina; Reserve Betampona, Camp Rendrirendry 34.1 km 332° Toamasina; F.C. Sandranantitra; F.C. Didy; F.C. Andriantantely; P.N. Mantadia; Analamay; Foret Ambatovy, 14.3 km 57° Moramanga; Torotorofotsy; Andasibe National Park, botanic garden near eIntrance, West of ANGAP office; Res. Analamazaotra, Parc National, Andasibe; Fianarantsoa: Foret d'Atsirakambiaty, 7.6 km 285° WNW Itremo; Ranomafana Nat. Park, Miaranony Forest; Vohiparara broken bridge, Fianarantsoa Prov. ; Parc National de Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana; Parc Nationale Ranomafana: Talatakely; 3 km W Ranomafana, nr. Ifandiana; research cabin at Talatakely, Ranomafana National Park ; radio tower, Ranomafana National Park, Fianarantsoa Prov. ; Namorona River at footbridge, Ranomafana National Park; Ranomafana National Park, Tavolo tree; Belle Vue trail, Ranomafana National Park, Fianarantsoa Prov. ; 7 km W Ranomafana; Vatoharanana; Parc National de Ranomafana, Vatoharanana River, 4.1 km 231 ° SW Ranomafana; P.N. Ranomafana, Vatoharanana-Ankovoka; 8 km E Kianjavato, Vatovavy Forest; 7.6 km 122° Kianjavato, Foret Classee Vatovavy; 2 km W Andrambovato, along river Tatamaly; Foret d'Ambalagoavy Nord, Ikongo, Ambatombe; 45 km S. Ambalavao; 45 km S Ambalavao; 43 km S Ambalavao, Res. Andringitra ; Parc National d'Isalo, Ambovo Springs, 29.3 km 4° N Ranohira; 8.0 km NE Ivohibe; 9.0 km NE Ivohibe; R.S. Ivohibe, 7.5 km ENE Ivohibe; Parc National d'Isalo, 9.1 km 354° N Ranohira; Foret d'Analalava, 29.6 km 280° W Ranohira; Foret de Vevembe, 66.6 km 293° Farafangana; Province Toliara: Reserve Speciale d'Ambohijanahary, Foret d'Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala; Reserve Speciale d'Ambohijanahary, Foret d'Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala; Vohibasia Forest, 59 km NE Sakaraha; southern Isoky-Vohimena Forest, 59 km NE Sakaraha; Foret Classee d'Analavelona, 33.2 km 344° NNW Mahaboboka; Foret Classee d'Analavelona, 29.2 km 343° NNW Mahaboboka; Foret Classee d'Analavelona, 29.4 km 343° NNW Mahaboboka; Foret de Tsinjoriaky, 6.2 km 84° E Tsifota; Parc National de Zombitse, 19.8 km 84° E Sakaraha; Parc National de Zombitse, 17.7 km 98° E Sakaraha; 15 km E Sakaraha; Foret de Mite, 20.7 km 29° WNW Tongobory; Sept Lacs; Beza-Mahafaly, 27 km E Betioky; Ehazoara Canyon, 26 km E Betioky; 70.7 km NNE Tolanaro, Mahermano Mt.; 11 km NW Enakara, Res. Andohahela; 10 km NW Enakara, Res. Andohahela; Res. Andohahela, 6 km SSW Eminiminy; Parc National d'Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro; Parc National d'Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro; 2.7 km WNW 302° Ste. Luce; 9.2 km N Tolanaro, Ilapany Mt . ; 29.5 km WNW Tolanaro, Vasiha Mt. ; Parc National d'Andohahela, Foret d'Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro; Mandena, 8.4 km NNE 30° Tolagnaro; Reserve Prive Berenty, Foret de Bealoka, Mandrare River, 14.6 km 329° NNW Amboasary; Reserve Prive Berenty, Foret de Malaza, Mandrare River, 8.6 km 314° NW Amboasary; Reserve Berenty; Foret de Petriky, 12.5 km W 272° Tolagnaro; 4.4 km 148° SSE Lavanono; Reserve Speciale de Cap Sainte Marie, 14.9 km 261° W Marovato; near road, Zombitse National Park, Tulear Prov.; near ANGAP office, Zombitse National Park, Tulear Prov.; Parcel I, Beza Mahafaly Reserve, near research station, Tulear Province; Tsimelahy - Parcel II, Andohahela National Park, transition forest, Tulear Province.