Pheidole megacephala (Fabricius)

(Figs. 21A–F)

Formica edax Forskål, 1775: 84 (nomen oblitum).

Syntype (s): (minor and major worker(s)?): EGYPT [not examined]. Junior synonym of Pheidole megacephala: Dalla Torre, 1892: 90; Emery, 1892: 160.

Formica megacephala Fabricius, 1793: 361 .

NEOTYPE (major worker) by present designation: MAURITIUS, Camizard Mt., Bambous, 20.3328 S, 57.723 E, 375 m, rainforest, ex rotten log, collection code BLF12051, 27.v.2005 (B.L. Fisher et al.) (CASC: CASENT0104990). [previous syntype (s), from MAURITIUS (‘Ile de France’), presumed lost]. Description of queen: Latreille, 1802: 232; descriptions of minor worker, major worker, queen, male: Mayr, 1861: 70; descriptions of larva: Wheeler, G.C. & Wheeler, J. 1953: 75. Combination in Pheidole: Roger, 1863: 30 .

Myrmica trinodis Losana, 1834: 327 .

Syntype (s) (‘worker’): ITALY [not examined]. Junior synonym of Pheidole megacephala: Roger, 1863: 30 .

Oecophthora pusilla Heer, 1852: 15 .

Syntype (s) (minor and major worker(s), queen(s), male(s)): PORTUGAL [not examined]. Combination in Pheidole: Smith, F. 1858: 173; subspecies of Pheidole megacephala: Emery, 1915: 235; senior synonym of Pheidole laevigata Smith: Roger, 1859: 259; subspecies of Pheidole megacephala Emery, 1915: 235; senior synonym of Pheidole laevigata Mayr: Mayr, 1870: 981; junior synonym of Pheidole megacephala: Wheeler, W.M. 1922: 812 .

Pheidole janus Smith, 1858: 175 .

Syntype (s) (minor and major worker(s)): CEYLON [not examined]. Junior synonym of Pheidole pusilla: Mayr, 1886: 360 . Junior synonym of Pheidole megacephala: Wheeler, W.M. 1922: 812 .

Myrmica(?) laevigata Smith, 1855: 130 .

Syntypes (s) (minor worker(s)): GREAT BRITAIN [not examined]. Junior synonym of Pheidole pusilla: Roger, 1859: 259; junior synonym of Pheidole pallidula: Smith, 1858 b: 282; junior synonym of Pheidole megacephala: Roger, 1863: 30; junior synonym of Pheidole pusilla: Emery, 1915: 235 . Junior synonym of Pheidole megacephala: Wheeler, W.M. 1922: 812 .

Myrmica agilis Smith, F., 1857: 71 .

Syntype (s): (minor worker(s)?): WEST MALAYSIA [not examined]. Combination in Pheidole: Donisthorpe, 1932: 449; junior synonym of Pheidole megacephala: Eguchi, 2008: 56 .

Pheidole laevigata Mayr, 1862: 747 .

Syntype (s): (major worker (s)): BRAZIL [not examined]. Unresolved junior secondary homonym of Pheidole laevigata Smith, F. Junior synonym of Pheidole pusilla: Mayr, 1870: 981 . Junior synonym of Pheidole megacephala: Wheeler, W.M. 1922: 812 .

Atta testacea Smith, F. 1858: 168 .

Syntype (s) (minor and major worker(s)): BRAZIL [not examined]. Combination in Pheidole: Mayr, 1886: 360; junior synonym of Pheidole megacephala: Brown, 1981: 530 .

Oecophthora perniciosa Gerstäcker, 1859: 263 .

Syntype (minor worker): MOZAMBIQUE [examined]. Combination in Pheidole: Roger, 1863: 30; junior synonym of Pheidole megacephala: Emery, 1915: 235 (ZMHB) .

Myrmica suspiciosa Smith, F., 1859: 148 .

Syntype (s) (minor worker(s)?): INDONESIA (Aru Is.) [not examined]. Junior synonym of Pheidole megacephala: Donisthorpe, 1932: 455 .

Pheidole megacephala var. scabrior Forel, 1891: 178 (See also: Forel, 1897: 188).

Syntypes (2 minor workers, 1 major worker, 3 queens, 1 male): MADAGASCAR, Nosibé ( Voeltzkow) (MHNG) [examined]. Syn. n.

Pheidole picata Forel, 1891: 178 .

Syntypes (2 major workers, 3 minor workers, 2 queens, 1 male): MADAGASCAR, Antananarivo ( Camboué) (MHNG) [examined]. (Variation of megacephala: Forel, 1895: 49; subspecies of punctulata: Forel, 1897: 186; Forel, 1905: 163; Santschi, 1910: 370; raised to species: Emery, 1915: 245; Wheeler, W.M. 1922: 1019). Syn. n.

Pheidole punctulata r. gietleni Forel, 1905: 164 .

Syntypes (2 major workers, 2 minor workers): MADAGASCAR, Fianarantsoa ( Gietlen) (MHNG) [examined]. Subspecies of Pheidole picata: Emery, 1915: 245 . Syn. n.

Pheidole picata var. bernhardae Emery, 1915: 245 (First available use of Pheidole punctulata spinosa bernhardae Forel, 1905: 164; unavailable name).

Syntypes (1 major worker, 1 minor worker): MADAGASCAR, Fianarantsoa ( Gietlen) (MHNG) [examined]. Syn. n.

Workers associated with the neotype: (4 major & 18 minor workers) same data as neotype (+ on low vegetation; ground foragers), collection codes BLF12047, BLF12051, BLF12053, BLF12055, BLF12059 (CASC: CASENT0055882 – CASENT0055886, CASENT0055903 – CASENT0055905, CASENT0059653 – CASENT0059657, CASENT0059671 – CASENT0059674; SAMC: CASENT0059652); (3 major & 16 minor workers), MAURITIUS, Moka Range, Le Pouce Mt., 20.1985 S, 57.529 E, 750 m, closed vegetation, under stone / ex rotten log / under tree bark, live tree, collection codes BLF12002, BLF12003, BLF12007, 25.v.2005 (B.L. Fisher et al.) (BMNH: CASENT0056010; CASC: CASENT0056004 – CASENT0056009, CASENT0056011 – CASENT0056019); (4 major & 17 minor workers), MAURITIUS, Moka Range, Pieter Both Mt., 20.1922 S, 57.5553 E, 770 m, closed vegetation, ground nest, collection codes BLF12028, BLF12035, BLF12036, BLF12037, 26.v.2005 (B. L. Fisher al.) (CASC: CASENT0055944 – CASENT0055947, CASENT0055951 – CASENT0055957, CASENT0059592 – CASENT0059594, CASENT0059597; MCZC: CASENT0059595, MHNG: CASENT0059596) .

Diagnosis: Moderately small species (WL major 0.99–1.22 mm, WL minor 0.62–0.84 mm), with moderately short scapes (SI minor 114–122, major SI 47–58) and legs (FI minor 114–125, FI major 65–73), both workers subcastes with acute short-spinose propodeal spines, absent promesonotal process, and postpetiole with large convex ventral process, mesosoma with relatively few, usually stiff and often blunt or apically bifurcate standing hairs, scape pilosity subdecumbent to suberect, hind tibia pilosity on inner edge decumbent, along outer edge subdecumbent to suberect. Major head often heart shaped, with moderately deep posterior emargination, shortirregular longitudinal rugae anteriorly and sculpture absent from posterior 2/5 of head, except for two weak to superficial longitudinal rugulae medially, hypostomal margin without median process and very small to inconspicuous submedian teeth, promesonotum almost completely smooth and shiny, remainder of mesosoma weakly punctate, smooth or superficially sculptured spots usually present on both katepisternum and lateropropodeum, postpetiole on average 1.9 times wider than petiole (PpWI 171–210). Minor head shape usually ovoid and posterior head margin slightly compressed, head almost as wide as long (CI 86–92), face completely smooth and shiny, except for sometimes weakly developed concentric rugulae around antennal insertions, postpetiole on average 0.8 times as long as petiole (PpLI 70–91).

Description of major workers: Measurements (neotype): HW 1.29, HL 1.28, SL 0.69, MDL 0.67, EL 0.17, WL 1.04, PNH 0.45, PNW 0.60, MNH 0.45, PDH 0.34, PTL 0.38, PPL 0.26, PTH 0.22, PPH 0.26, PTW 0.18, PPW 0.36, PSL 0.15, MFL 0.92, MTL 0.66, CI 101, SI 54, MDI 52, EI 13, FI 71, PSLI 12, LPpI 100, DPpI 137, PpWI 197, PpLI 68, PpHI 118.

Measurements (n=19): HW 1.10–1.54 (1.30), HL 1.04–1.59 (1.30), SL 0.59–0.76 (0.67), MDL 0.59–0.82 (0.70), EL 0.15–0.19 (0.17), WL 0.94–1.22 (1.06), PNH 0.36–0.54 (0.45), PNW 0.54–0.74 (0.62), MNH 0.61–0.80 (0.68), PDH 0.33–0.53 (0.37), PTL 0.31–0.44 (0.38), PPL 0.23–0.29 (0.26), PTH 0.20–0.28 (0.23), PPH 0.22–0.31 (0.26), PTW 0.14–0.22 (0.18), PPW 0.27–0.45 (0.34), PSL 0.13–0.20 (0.16), MFL 0.77–1.06 (0.89), MTL 0.59– 0.77 (0.67), CI 97–106 (101), SI 47–58 (52), MDI 51–57 (54), EI 11–15 (13), FI 65–73 (69), PSLI 11–14 (12), LPpI 90–114 (100), DPpI 108–156 (134), PpWI 171–210 (190), PpLI 61–81 (69), PpHI 105–123 (114).

Head as long as wide (CI 97–103), usually heart shaped with strongly convex sides and posterior emargination moderately deeply impressed. Mandibles moderately long (MDI 51–57), smooth and shiny. Clypeus smooth with median carina short or absent and one or two pairs of lateral carinae. Frons and sides of head anteriorly, longitudinally rugose, interspaces superficially punctate at frons, punctate or weakly punctate at sides, with a few longer rugae ventral of eyes, posterior 2/5 of head smooth and shiny, except for two weakly to faintly developed longitudinal rugae medially and, rarely present, short, faint rugulae submedially at posterior emargination. Sides of head dorsal and ventral of eyes in profile with few irregular rugae extending up to posterior 1/4 of head, space in between, posterior of eyes, largely unsculptured. Frontal carinae and antennal scrobe absent. Scapes of medium length (SI 47–58) with decumbent to subdecumbent pilosity and few longer, suberect hairs along outer edge. Submedian hypostomal teeth very small to inconspicuous, median process absent. Promesonotum moderately high-domed, in profile usually subangulate posteriorly, humeri in dorsal view not produced, promesonotal process not conspicuously produced, posterior declivity often weakly marginate anteriorly (at posterior end of pronotum), oblique and angulate at its lower portion. Surface of promesonotum and lateropronotum mostly smooth to superficially punctate, with few faint, irregular, transverse rugulae, remainder of mesosoma weakly punctate, with smooth or superficially punctate spots between. Metanotal groove narrow, barely or not impressed, and with few superficially developed cross-ribs. Spines short-spinose to subtriangular in profile (PSI 11–14). Metatibia relatively short (FI 65–72), metafemur pilosity on inner edge decumbent, on outer edge suberect to subdecumbent. Postpetiole in profile about as high as long (LPpI 90–114), with large convex ventral process, in dorsal view trapezoid and on average 1.9 times wider than petiole (PpWI 171–210). Dorsum of petiole and postpetiole smooth, remainder weakly punctate. Gaster smooth. Amount and length of standing hairs on body dorsum variable, usually with few long hairs and some shorter hairs, with several hairs apically blunt or bifurcate. Fine, short pilosity not abundant. Color variable from light to dark brown, head and gaster often darker than the rest.

Description of minor workers: Measurements (n=20): HW 0.50–0.61 (0.55), HL 0.57–0.68 (0.62), SL 0.61– 0.72 (0.65), MDL 0.32–0.41 (0.36), EL 0.12–0.15 (0.13), WL 0.62–0.84 (0.73), PNH 0.22–0.29 (0.25), PNW 0.32–0.39 (0.35), MNH 0.42–0.53 (0.46), PDH 0.21–0.27 (0.24), PTL 0.22–0.28 (0.24), PPL 0.17–0.22 (0.19), PTH 0.13–0.16 (0.14), PPH 0.16–0.19 (0.14), PTW 0.09–0.11 (0.10), PPW 0.17–0.21 (0.18), PSL 0.07–0.10 (0.08), MFL 0.57–0.76 (0.67), MTL 0.44–0.57 (0.51), CI 86–92 (90), SI 114–122 (118), MDI 63–70 (66), EI 22– 26 (24), FI 114–125 (121), PSLI 13–16 (14), LPpI 94–120 (109), DPpI 89–112 (98), PpWI 164–200 (180), PpLI 70–91 (79), PpHI 110–146 (125).

Head shape roundly ovoid, slightly longer than wide (CI 86–92), sides convex, posterior head margin rounded laterally, medially sometimes slightly compressed, occipital carina very narrow in full-face view, often weakly impressed medially. Mandibles relatively short (MDI 63–70), smooth. Clypeus and remainder of face smooth, clypeal carinae absent or inconspicuous, around antennal insertion few concentric rugulae present and often weakly to superficially developed. Scapes relatively short (SI 114–122), when laid back surpassing posterior head margin by slightly more than the length of tenth funicular segment, with subdecumbent to suberect pilosity. Promesonotal outline in lateral view roundly convex, usually evenly declining from highest point toward metanotal groove, posterior process absent. Metanotal groove in profile weakly impressed, with dorsally weak to laterally superficial cross-ribs. Propodeum about as long as high, in profile slightly declining toward spines. Propodeal spines much shorter than distance between their bases, short-spinose to subtriangular and acute (PSLI 13–16, mean: 14). Promesonotum largely smooth, except superficial punctures anteriorly near the neck, mesopleuron and propodeum punctate to weakly punctate, often with superficially sculptured to smooth areas laterally. Metafemur short (FI 114–125), metatibia with decumbent pilosity and scattered suberect longer hairs on outer edge. Postpetiole in profile with large convex ventral process, on average 1.1 times longer than high (LPpI 94–120), and 0.8 times as long as petiole (PpLI 70–91). Dorsum of petiole and postpetiole smooth, remainder weakly to superficially punctate. Gaster smooth and shiny. Standing hairs short to moderately long, relatively coarse, erect to suberect, not abundant, and usually with some to several hairs apically split or bifurcate. Color usually light brown, sometimes brown, gaster darker brown.

Discussion: Pheidole megacephala – commonly also known as the ‘African big–headed ant’ – is a globally distributed invasive ant species and can be found on almost every island in the Malagasy region. This species is considered one of the most problematic invasive ant species and has caused devastating environmental problems all over the world (Hoffmann et al. 1999, Hoffmann & Parr 2008, Wetterer 2012). First described by Fabricius from the former ‘Ile de France’, called Mauritius today, the origins of this species remain unknown. Pheidole megacephala and the megacephala group are probably native to the Afrotropics (Wetterer 2012), although a Malagasy origin is also a possibility, as the whole group seems to be quite diverse in Madagascar.

Possibly as a result of its wide distribution and local abundances, a number of subspecies and synonyms have accumulated during the nineteenth and early twentieth century. Several of the subspecies are probably synonyms (unpublished data), and a number of described and undescribed related species add to the taxonomic confusion within the megacephala group. The types are presumed lost. No type specimens could be located in both of the two possible collections – the Zoological Museum of the University of Copenhagen (ZMUC) and the Muséum National d'Histoire Naturelle in Paris (MNHN). To stabilize the name P. megacephala we designated a neotype in this publication. The neotype, together with many associated specimens, was collected on Mauritius. The material is morphologically concordant with previous redescriptions (Wilson 2003, Eguchi 2008), and identical to P. megacephala material from other collection localities (e.g. Papua New Guinea and Australia).

Morphologically, both the minor and major workers of P. megacephala display relatively few diagnostic characters to set them apart from other members of the group. On the Comoros P. megacephala also occurs together with the two newly described species P. decepticon and P. megatron . The minor workers of the latter two species are very similar in morphometric measurements and external morphology to those of P. megacephala, and single major workers possess few clearly defined features that can be used for definite species identifications (see also in respective discussions). Because P. megacephala is a morphologically variable species, series with several major workers are much more helpful for positive identifications. Especially the majors of P. megacephala and P. decepticon are in many characters almost identical and both vary significantly in their development of the typically heart-shaped head. The most apparent and stable differences, however, seem to be the amount of sculpture on the posterior half of the major worker’s head, the type and overall quantity of standing hairs on the meso- and metasoma, as well as the shape of the postpetiolar ventral processes. Further minor differences in the morphologies of P. decepticon suggest ongoing differentiation between populations on the islands where it occurs.

Additional material examined: AUSTRALIA: Norfolk, No. 3992, 28.xii.37; 01.iv.1938; Queensland, Parish of Bunker, Great Barrier Reef, Heron Island, 26-30.vii.1984 (T.W. Davies); BELIZE: South Water Cay, 01.vi.1985 (J.F. Lynch); COMOROS: Anjouan: Moya village, S 8.23148w, 10m, suburban area, on stone wall near sea, 04.xii.1994 (Roger); KENYA: Tsavo Park, Ati R., 26.vi.1967 (G. Sibley); MADAGASCAR: Antsiranana: R.S. Manongarivo, 10.8 km 229° SW Antanambao, 13.9617 S, 48.4333 E, 400 m, rainforest, 11.viii.1998 (B.L. Fisher); Toamasina: Antaratasy, 43.5 km 23° Toamasina, 17.7673 S, 49.4777 E, 15 m, urban garden, 02.viii.2006 (B. Blaimer, F.N. Raharimalala); Ampasimbe, 118 km 17° Toamasina, 17.0928 S, 49.4778 E, 25 m, urban garden, 07.viii.2006 (B. Blaimer, F.N. Raharimalala); Antetezambaro, 16.0 km 21° Toamasina, 17.0528 S, 49.567 E, 10 m, urban habitats, 08.i.2006 (B. Blaimer, F.N. Raharimalala); Mahambo, 73.8 km 16° Toamasina, 17.4893 S, 49.4517 E, 10 m, urban garden, 04.viii.2006 (B. Blaimer, F.N. Raharimalala); Toamasina-Ville, 18.1552 S, 49.4095 E, 10 m, interior house, 31.vii.2006 (B. Blaimer, F.N. Raharimalala); Fianarantsoa: Farafangana, 22.8197 S, 47.83 E, 10 m, urban gardens, 25.iv.2006 (B.L. Fisher et al.); MAURITIUS: 16.viii.2004, trail across jeep track (L. Lach); 20.315 S, 57.4469 E, 231 m, Dry II forest, 27.iv.2009 (J. Casquet); Basin Blanc, 20.4532 S, 57.4777 E, 480 m, rainforest, 29.v.2005 (B.L. Fisher et al.); Belle Rive, L'Etoile, 17.xii.2006 (M. Madl); Black River, Corps de Garde, 20.2633 S, 57.4549 E, 684 m, 24.vii.2004 (L. Lach); Black River, Florin, 20.3911 S, 57.4611 E, 631 m, 29.xi.2004 (L. Lach); Black River, Macchabee, 20.4005 S, 57.4552 E, 656 m, 22.xi.2004 / 04.iv.2005 / 27.vi.2005 (L. Lach); Black River, Petrin, 20.4063 S, 57.4666 E, 644 m, 25.v.2005 (A.V. Suarez); Blue Bay, 20.4414 S, 57.7192 E, 20.x.2004 (L. Lach); Gunners Quoin (N. Cole); Moka Range, Le Pouce, 20.2 S, 57.5167 E, 700 m, grassland, 09.iv.1989 (P.S. Ward); Moka Range, Le Pouce, 26.ix.2004 (L. Lach); Side of Le Pouce, 20.1906 S, 57.5058 E, 173 m, 1/2 dry II forest, 19.iv.2009 (J. Casquet); Side of Le Pouce, 20.1917 S, 57.5067 E, 176 m, 100% butterfly liana, 19.iv.2009 (J. Casquet); Moka Range, Perrier, 20.3521 S, 57.4942 E, 564 m, 12.xi.2004 (L. Lach); Moka Range, Petrin, 20.4063 S, 57.4666 E, 644 m, 29.xi.2004 / 17.iii.2005 / 22.vi.2005 (L. Lach); La Nicolière Reserve, 06.v.2007 (M. Madl); Macchabee Forest, 20.4 S, 57.45 E, 600 m, disturbed rainforest, 10.iv.1989 (P.S. Ward); Macchabee View Point, 20.x.2004 (L. Lach); Montagne du lion, 20.3631 S, 57.7261 E, 393 m, wet II vegetation, 25.iv. 2009 (J. Casquet); On the trail to Piton Savane, 20.4419 S, 57.4875 E, 675 m, forest road surrounded by wet II forest, 16.iv.2009 (J. Casquet); Perrier Nature Reserve, 20.3521 S, 57.4942 E, 564 m, 25.v.2005 (A.V. Suarez); Petite Rivière Noire Mt., 20.4088 S, 57.4077 E, 750 m, rainforest, 29.v.2005 (B.L. Fisher et al.); Savanne, Bel Ombre-Belloughy, 20.4689 S, 57.4272 E, 82 m, 19.xi.2004 / 23.xi.2004 / 25.ii.2005 / 30.v.2005 / 14.vii.2005 / 02.viii.2005 (L. Lach); Savanne, Combo-lower, 20.4722 S, 57.5153, 320 m, 13.xii.2004 / 13.iv.2005 / 11.vii.2005 (L. Lach); St. Catherine (Linden); Yemen, 14.xii.2006 (M. Madl); Ile Aux Aigrettes: 03.ix.2004 / 06.ix.2004 (L. Lach); 20.419 S, 57.7302 E, on island, 23.v.2005 (A.V. Suarez); 20.4188 S, 57.7305 E, 1 m, coastal scrub, 28.v.2005 (B.L. Fisher et al.); 20.4197 S, 57.7311 E, 25 m, protected coastal forest, 21.iv.2009 (J. Casquet); Ile de la Passe: (N. Cole); Ile Marianne: 20.3667 S, 57.7833 E, 5 m, 02.iii.2005 (L. Lach); Rodrigues: Anse Quitor, 19.7561 S, 63.3667 E, 21 m, 15.vi.2005, (L. Lach); Canyon Tyeul, 19.7421S, 63.375 E, 35 m, degraded dry forest, 14.x.2007 (A. Meunier); Caverne Bambara, 19.7416 S, 63.3739 E, 38 m, degraded dry forest, 14.xi.2007 (A. Meunier); Citronelle, 19.7149 S, 63.4163 E, 10.xii.2007 (M. Madl); Cotton Bay Hotel, 19.7149 S, 63.4163 E, 25.iv.2008 (M. Madl); Montagne des 4 vents, 19.7181 S, 63.4117 E, 303 m, Low & dry II forest, 02.v.2009 (J. Casquet); Round Island: 19.8508 S, 57.7697 E, 206 m, 01.viii.2004 (R. Bone); 18.viii.2004 (R. Bone); 19.8512 S, 57.7838 E, 102 m, 21.i.2005 (L. Lach); 19.8508 S, 57.7697 E, 206 m, 22.i.2005 (L. Lach); 19.8471 S, 57.789 E, 296 m, 23.i.2005 (L. Lach); 19.8568 S, 57.7873 E, 82 m, 23.i.2005 (L. Lach); 23.i.2005 (L. Lach); 24.i.2005 (L. Lach); MAYOTTE: Benara, Labattoir, 12.793 S, 45.2761 E, 60 m, diverse vegetation in understory of mango and coconut plantations, 16.i.2010 (J. Rochat); Baie de Tsingoni, 12.7926 S, 45.1076 E, 5 m, mangrove, coastal scrub, 26.xi.2007 (B.L. Fisher et al.); Coconi DAF campus, dense wet forest, 22.vii.1998 (R. Jocque); Combani, 12.8107 S, 45.2793 E, 7 m, native littoral and secondary vegetation, 15.i.2010 (G. Paulus / S. Gasnier); Combani, 12.8134 S, 45.2813 E, 24 m, native littoral and secondary vegetation, 16.i.2010 (J. Rochat); Mont Combani, 12.8063 S, 45.1531 E, 370 m, rainforest, 25.xi.2007 (B.L. Fisher et al.); Dapani, 12.9749 S, 45.1618 E, 1 m, mangrove, coastal scrub, 27.xi.2007 (B.L. Fisher et al.); Sazile, 12.9784 S, 45.1726 E, 35 m, dry forest, 27.xi.2007 (B.L. Fisher et al.); Tanaraki, 12.7575 S, 45.0678 E, 10 m, coastal scrub, 26.xi.2007 (B.L. Fisher et al.); PAPUA NEW GUINEA: Morobe Bululo: (53 km W), Aseki Rd., 7º19'22" S, 146º21'20" E, 2100 m, mixed montane forest litter, 09.ii.2000 (R.S. Anderson); Morobe Wau: (19 km NNW), 7º24'11" S, 146º39'41" E, 2150 m, montane forest litter, 06.ii.2000 (R.S. Anderson); (9 km NNW), Mt. Kaindi, 7º20'24" S, 146º40'01" E, 2050 m, montane forest litter, 06.ii.2000 (R.S. Anderson); (25 km SE), Kuper Range, 7º30'14" S, 146º48'13" E, 2200 m, montane forest litter, 07.ii.2000 (R.S. Anderson); RÉUNION: Basse Vallée, Théophane Bègue, guesthouse, 21.3481 S, 55.7075 E, 449 m, in garden, 21.iii.2009 (J. Casquet); Cilaos, Ilet des Salazes, 21.1116 S, 55.4493 E, 1575 m, mountain forest/garden, 31.iii.2011 (B.L. Fisher et al.); Etang Salé, dry forest, disturbed area, 24.iii.2009 (J. Casquet); Grande Chaloupe, 20.9293 S, 55.3888 E, 580 m, tropical dry forest, 20.v.2007 (B.L. Fisher et al.); Hermitage, 21.0855 S, 55.2275 E, 20 m, urban garden, 21.v.2007 (B.L. Fisher et al.); Mare Longue, 22.viii.2001 (F. Blard); Mare Longue, 21.3642 S, 55.7435 E, 30 m, rainforest, 22.v.2007 (B.L. Fisher et al.); Saint-Denis, Le Port, 20.9492 S, 55.3095 E, 70 m, garden, 20.vi.2002 (F. Blard); Saint Denis, 20.8862 S, 55.4847 E, 10 m, urban garden, 21.v.2007 (B.L. Fisher et al.); S EYCHELLES: Aldabra Island: Grande Terre, Takamaka grove, 15.xii.2008 (V. Robert); Malabar: Camp Malabar 15.xii.2005 (J. Gerlach); Aride Island: 01.vii.2000 (J. Gerlach); 03.xii.2008 (V. Robert); 4.21273 S, 55.6645 E, P. grandis forest 01.vii.2010 (R. Gaigher); 4.21407 S, 55.6682 E, 10 m, native forest, 05.ii.2005 (B.L. Fisher et al.); Assumption Island: Airport, 19.xii.2008 (V. Robert); Bijoutier Island (Alphonse): 08.iv.2001 (J. Gerlach); Bird Island: 01.x.2010 (J. Gerlach); Cosmoledo Atoll (Wizard Island): 9.73528 S, 47.6478 E / 9.7425 S, 47.6517 E / 9.73833 S, 47.6542 E / 9.74 S, 47.6533 E / 9.76028 S, 47.6419 E / 9.75111 S, 47.6494 E / 9.75444 S, 47.6481 E / 9.75306 S, 47.6489 E, 10 m, lowland coastal, coralline island, 16. & 21.xi.2007 (G. Galman); Grande Ile, 17.xii.2005 (J. Gerlach); Menai, 17.xii.2005 (J. Gerlach); Cousine Island: 4.3507 S, 55.6478 E, P. grandis forest 01.vii.2010 (R. Gaigher); 4.35009 S, 55.6468 E, 60 m, Ficus forest on rocks, 07.iii.2008 (B.L. Fisher); Félicité Island: 4.32012 S, 55.8668 E, 20 m, coastal scrub, 01.ii.2010 (B.L. Fisher et al.); La Digue Island: La Passe, & La Veuve, 06.xii.2008 (V. Robert); Mahé Island: Cascade, 4.67494 S, 55.4982 E, 150 m, roadside, 09.ii.2010 (B.L. Fisher et al.); Mont Copolia 4.65121 S, 55.4584 E, 520 m, forest, 08.ii.2010 (B.L. Fisher et al.); Montagne, Posée, Palm Forest Reserve, Brulée, 4.712 S, 55.5019 E, 360 m, palm forest, 08.iii.2008 (B.L. Fisher); Morne Blanc, 4.65988 S, 55.4374 E, 480 m, mixed forest near road, 10.ii.2010 (B.L. Fisher et al.); North Island: 30.vii.2000 (J. Gerlach); Praslin Island: Grande Anse, 05.xii.2008 (V. Robert); Silhouette Island: Above Jardin Marron on crest to Mont Plaisir and Pot à Eau, 4.4867 S, 55.2341 E, 520 m, forest, 20.i.2010 (B.L. Fisher et al.); La Passe, 4.48484 S, 55.2507 E, 35 m, park/garden, 23.i.2010 (B.L. Fisher et al.); on coastal path to Anse Mondon, 4.46951 S, 55.2416 E, 10 m, coastal scrub, 23.i.2010 (B.L. Fisher et al.); TANZANIA: Unguga region, Josani For. Res. Zanzibar, Ground Water Forest, 6.2676 S, 39.4111 E, 19 m, 30.x.-14.xi.2007 (P. Hawkes, M. Bhoke, U. Richard); Lindi region, Mbarawala Plateaux, Primary Forest, 9.0396 S, 39.1201 E, 270 m, 29.ii.- 04.iii.2008 (P. Hawkes, Y. Mlacha, F. Ninga); Morogoro region, Mamiwa-Kizara For. Res., Primary Forest, 6.3753 S, 36.9371 E, 1989 m, 16.-21.viii.2005 (P. Hawkes, J. Makawati, R. Mtana); UGANDA: Entebbe, Botanical Garden, 0.0602 N, 32.4807 E, 1130 m, 19.viii.2012 (G. Fischer); USA: Phoenix, Arizona, 13.x.1980 (R.R. Snelling?).