Pheidole fervens

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Pheidole fervens
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Attini
Genus: Pheidole
Species group: fervens
Species: P. fervens
Binomial name
Pheidole fervens
Smith, F., 1858

Pheidole fervens casent0008638 profile 1.jpg

Pheidole fervens casent0008638 dorsal 1.jpg

Specimen Label

Subspecies
Synonyms


Common Name
Minami-oozu-ari
Language: Japanese

Fischer & Fisher (2013): Pheidole fervens, described from Singapore, is a widespread invasive species and could be native to the Oriental or the Oceanic region (Eguchi 2004, Wilson & Taylor 1967). It usually nests in soil or under stones and usually prefers disturbed habitats (Eguchi 2004). In Japan it is found nesting under stones in open land grading to forest margins (Ogata, 1981). On Fiji it was collected in elevations between 1–800 m and tended to inhabit several different habitats from forest edge and mangrove forest to disturbed and undisturbed rainforest. On Mauritius, which seems to be its western distribution limit, this ant was found in the leaf litter of lowland rainforest (200 m elevation). As with most other introduced ants, such as Pheidole indica, it is unclear what effect, if any, P. fervens might have on the rest of the local ant fauna. Considering that the ecosystems of the Mauritius islands have been severely altered and disturbed since the arrival of human settlers several hundred years ago, and native species must also contend with the invasions of introduced organisms (Fisher 2005, Ward 1990), parts of the original ant fauna might have been marginalized or driven to extinction long ago. We speculate that the scarcity of P. fervens specimens in Mauritius ant collections and its presence in the rainforest means that this ant has not yet become an invasive and dominant aggressor toward other ant species. Nevertheless, efforts to further investigate the presence and activity of introduced Pheidole species are worthwhile, especially in the few remaining undisturbed habitats.

Pashaei Rad et al. (2018) found this species in Iran on the ground in moist to moderate rainfall areas.

At a Glance • Invasive  

Identification

Fischer and Fisher (2013) - Large species (WL major 1.06–1.17, WL minor 0.82–0.91), minor workers with relatively long scapes and legs (SI 133–154, FI 142–158) and major workers with long scapes and relatively long legs (SI 61–71, FI 84–91), both with promesonotal processes well developed and postpetiolar ventral process inconspicuous or very shallow. Major with frontal carinae reaching at least 4/5 of the distance to the posterior head margin, face longitudinally rugose, scrobe area and interspaces at sides of head punctate to weakly punctate, head in full-face view with coarse suberect hairs, eyes moderately large (EI 13–15), in profile metanotal groove vestigial to absent, propodeum sloped down toward spines, postpetiole in dorsal view trapezoidal and almost twice as wide as petiole (PpWI 172-195). Minor head oval and rounded posteriorly, occipital carina visible in full-face view but not well developed, face mostly smooth, often with fine, long rugulae at the sides, sometimes together with superficial punctures. Eyes moderately large (EI 21–25), dorsal promesonotum smooth, promesonotal process developed, subangulate in profile, metanotal groove well developed, impressed, propodeum relatively high and sloped toward spines.

Southwest Indian Ocean

Fischer and Fisher (2013) - The workers of P. fervens can be easily confused with those of Pheidole indica. Morphologically the minor workers are best separated by coarse, mostly suberect scape pilosity, impressed metanotal groove, relatively smaller eyes and shorter spines (EI 21–25, PSLI 10–13) in P. fervens versus decumbent to subdecumbent scape pilosity with longer suberect hairs along outer edge, inconspicuous metanotal groove in profile, and larger eyes and longer spines (EI 25–30, PSLI 14–19) in P. indica. The major workers of P. fervens are recognizable by having longer scapes (SI 61–71 versus SI 49–62) and a narrower postpetiole in dorsal view than P. indica (DPpI 133–150 and PpWI 172–195 versus DPpI 146–235 and PpWI 190–247). In P. fervens, the pilosity in the face and on the metatibia is coarser and at least partly suberect, and the punctures on the sides of the face are stronger than the thinner and more decumbent to subdecumbent pilosity and superficial punctures on the sides of the face in P. indica.

Indo-China

Eguchi (2008) - This medium-sized species with general habitus is similar to Pheidole binghamii, Pheidole elongicephala, Pheidole indica, Pheidole ochracea and Pheidole plagiaria among Indo-Chinese species. P. fervens is well separated from P. indica which has the following characteristics: eye relatively large (maximal diameter of eye much longer than antennal segment X in the major, and as long as or a little longer than X in the minor); in the major propodeal spine is relatively broadly based (see Eguchi 2004).

Posterior margin of head in full-face view is more deeply and narrowly concave in the major of P. binghamii, P. elongicephala and P. ochracea than in that of P. fervens, and sculpture on dorsum of vertexal lobe is usually stronger in the major of the former three.

The major of P. fervens is separated from that of Pheidole plagiaria which has the following characteristics: posterior margin of head in full-face view deeply concave; the rugulae running almost transversely along posterior margin of vertexal lobe.

Japan

Habitat preference, body size and coloration are similar to Pheidole indica, Pheidole noda and Pheidole megacephala. In fact, these species are sometimes closely sympatric in the Ryukyus. P. fervens can be distinguished by its lighter body color, smaller eyes, the relatively fine reticulation on the heads of its soldiers, the shape of the propodeal spines and the proportions of its postpetiole (Japanese Ant Image Database).

Keys including this Species

Distribution

Known from N. Vietnam, Manchurian subregion (southern part only), Oriental region, Austro-Malayan subregion, W. Pacific and West coast of N. America. In the Indo-Malayan subregion this species is one of the prevailing Pheidole species in semiurban and rural areas. (Eguchi 2008)

Latitudinal Distribution Pattern

Latitudinal Range: 32.812778° to -3.7°.

   
North
Temperate
North
Subtropical
Tropical South
Subtropical
South
Temperate

Distribution based on Regional Taxon Lists

Australasian Region: New Caledonia.
Indo-Australian Region: Borneo, Cook Islands, Fiji, French Polynesia, Guam, Indonesia (type locality), Kiribati, Krakatau Islands, Malaysia, Marshall Islands, Micronesia (Federated States of), Northern Mariana Islands, Palau, Philippines, Pitcairn, Samoa (type locality), Singapore (type locality), Tokelau, Tonga, Vanuatu.
Malagasy Region: Mauritius.
Nearctic Region: United States.
Oriental Region: India (type locality), Sri Lanka, Taiwan (type locality), Thailand, Vietnam.
Palaearctic Region: China (type locality), Iran, Japan (type locality).

Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
pChart

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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Biology

Association with Other Organisms

Explore-icon.png Explore: Show all Associate data or Search these data. See also a list of all data tables or learn how data is managed.
  • This species is a mutualist for the aphid Aphis gossypii (a trophobiont) (Idechiil et al., 2007; Saddiqui et al., 2019).

Castes

  • Liu, C. et al. 2020. Ants of the Hengduan Mountains, Figure 87, Pheidole fervens.

Worker

Minor

Images from AntWeb

Pheidole fervens casent0005780 head 1.jpgPheidole fervens casent0005780 profile 1.jpgPheidole fervens casent0005780 dorsal 1.jpgPheidole fervens casent0005780 label 1.jpg
Worker. Specimen code casent0005780. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by UCDC, Davis, CA, USA.
Pheidole-fervensH5.jpgPheidole-fervensL3.2.jpgPheidole-fervensD3.2.jpgPheidole-fervensLabel.jpg
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Pheidole fervens casent0008639 profile 1.jpgPheidole fervens casent0008639 dorsal 1.jpgPheidole fervens casent0008639 label 1.jpg
Worker. Specimen code casent0008639. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by UCDC, Davis, CA, USA.

Major

MCZ-ENT00020659 Pheidole javana-hef.jpgMCZ-ENT00020659 Pheidole javana-hal.jpgMCZ-ENT00020659 Pheidole javana-had.jpgMCZ-ENT00020659 Pheidole javana-var.jpgPheidole fervens casent0008638 head 1.jpgPheidole fervens casent0008638 profile 1.jpgPheidole fervens casent0008638 dorsal 1.jpgPheidole fervens casent0008638 label 1.jpgPheidole fervens casent0005779 head 1.jpgPheidole fervens casent0005779 profile 1.jpgPheidole fervens casent0005779 dorsal 1.jpgPheidole fervens casent0005779 label 1.jpgMCZ-ENT00020659 Pheidole javana-hef.jpgMCZ-ENT00020659 Pheidole javana-hal.jpgMCZ-ENT00020659 Pheidole javana-had.jpgMCZ-ENT00020659 Pheidole javana-var.jpg
.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • fervens. Pheidole fervens Smith, F. 1858b: 176 (s.) SINGAPORE. Ogata, 1982: 197 (m.). Senior synonym of cavannae, javana, nigriscapa and material of the unavailable name tahitiana referred here: Wilson & Taylor, 1967: 45; senior synonym of desucta: Eguchi, 2001b: 53; of amia, azumai, dharmsalana, dolenda, pungens, soror: Eguchi, 2004: 197. Current subspecies: nominal plus pectinata.
  • pungens. Solenopsis pungens Smith, F. 1861b: 48 (s.w.) INDONESIA (Sulawesi).
    • Combination in Pheidologeton: Emery, 1924d: 213; Donisthorpe, 1932c: 469; Chapman & Capco, 1951: 160;
    • Combination in Carebara: Bolton, 1995b: 333 (error); Fischer, et al. 2014: 72 (error);
    • Combination in Pheidole: Bolton, 1995b: 328.
    • Status as species: Roger, 1863b: 32; Mayr, 1863: 454; Smith, F. 1871a: 333; Dalla Torre, 1893: 77; Emery, 1924d: 213; Donisthorpe, 1932c: 469; Chapman & Capco, 1951: 160; Bolton, 1995b: 328.
    • Junior synonym of fervens: Eguchi, 2004: 198; Eguchi, 2008: 24; Fischer & Fisher, 2013: 322; Sarnat, et al. 2015: 38.
  • javana. Pheidole javana Mayr, 1867a: 98 (s.w.) INDONESIA (Java). Viehmeyer, 1916a: 121 (q.m.). Junior synonym of fervens: Wilson & Taylor, 1967: 45.
  • cavannae. Pheidole cavannae Emery, 1887b: 464 (footnote) (s.) NEW CALEDONIA. Subspecies of oceanica: Emery, 1914f: 401. Junior synonym of fervens: Wilson & Taylor, 1967: 45.
  • dharmsalana. Pheidole javana var. dharmsalana Forel, 1902c: 184 (s.), 198 (w.) INDIA. [Also described as new by Forel, 1902f: 546.] Subspecies of fervens: Bolton, 1995b: 320. Junior synonym of fervens: Eguchi, 2004: 198.
  • amia. Pheidole amia Forel, 1912a: 60 (s.w.) TAIWAN. Junior synonym of fervens: Eguchi, 2004: 197.
  • dolenda. Pheidole javana var. dolenda Forel, 1912a: 60 (s.w.) TAIWAN. Subspecies of fervens: Bolton, 1995b: 320. Junior synonym of fervens: Eguchi, 2004: 198.
  • nigriscapa. Pheidole (Pheidole) oceanica var. nigriscapa Santschi, 1928a: 48, fig. 3 (s.w.) SAMOA. Junior synonym of fervens: Wilson & Taylor, 1967: 45.
  • desucta. Pheidole javana var. desucta Wheeler, W.M. 1929f: 2 (s.w.q.) CHINA. Subspecies of fervens: Bolton, 1995b: 320. Junior synonym of fervens: Eguchi, 2001b: 53.
  • soror. Pheidole javana st. soror Santschi, 1937h: 369, fig. 6 (s.w.) TAIWAN. Subspecies of fervens: Bolton, 1995b: 330. Junior synonym of fervens: Eguchi, 2004: 198.
  • azumai. Pheidole nodus st. azumai Santschi, 1941: 274, fig. 1 (s.w.) JAPAN. Junior synonym of fervens: Eguchi, 2004: 198.

Type Material

  • Pheidole fervens: Lectotype (designated by Fischer & Fisher, 2013: 322), major worker, Singapore, The Natural History Museum.
  • Pheidole fervens: Paralectotype (designated by Fischer & Fisher, 2013: 322), 1 minor worker, Singapore, The Natural History Museum.
  • Solenopsis pungens: Lectotype (designated by Eguchi, 2004), major worker, Sulawesi, Indonesia, Oxford University Museum of Natural History.
  • Pheidole javana: Lectotype (designated by Eguchi, 2004), major worker, Java, Indonesia.
  • Pheidole javana dharmsalana: Lectotype (designated by Eguchi, 2004), major worker, India.
  • Pheidole amia: Lectotype (designated by Eguchi, 2004), major worker, Taiwan.
  • Pheidole javana dolenda: Lectotype (designated by Eguchi, 2004), major worker, Taiwan.
  • Pheidole oceanica nigriscapa: Syntype, 1 major worker, 7 minor workers, Samoa.
  • Pheidole javana desucta: Lectotype (designated by Eguchi, 2001), major worker, China.
  • Pheidole javana soror: Lectotype (designated by Eguchi, 2004), major worker, Taiwan.
  • Pheidole nodus azumai: Lectotype (designated by Eguchi, 2004), major worker, Japan.

The following notes on F. Smith type specimens have been provided by Barry Bolton (details):

Solenopsis pungens

Two syntype major workers and one syntype minor worker (on separate cards) in Oxford University Museum of Natural History. Each major worker labelled “Menado” and with a Smith det. label “Solenopsis pungens Smith.” The minor worker with “Men. 18” and an attached note explaining that Pheidole pungens was placed in Solenopsis by an oversight. Each of the three cards has the letter “c” in the lower left corner.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

Eguchi 2001. Figure 16.

Eguchi (2008) - Major (n=5). — HL 1.31–1.56 mm; HW 1.22–1.44 mm; CI 92–97; SL 0.84–0.95 mm; SI 66–70; FL 1.12–1.27 mm; FI 88–94. Minor (n=5). — HL 0.65–0.73 mm; HW 0.54–0.60 mm; CI 81–84; SL 0.77–0.87 mm; SI 138–153; FL 0.81–0.93 mm; FI 149–163.

Head in lateral view not or hardly impressed on vertex; posterior concavity of head in full-face view usually shallow; frons and anterior part of vertex rugose longitudinally; posterior part of vertex and dorsal face of vertexal lobe rugoso-reticulate, with interspaces weakly punctured; frontal carina conspicuous; antennal scrobe inconspicuous or shallowly impressed; clypeus without a median longitudinal carina; median and submedian processes of hypostoma absent or present but inconspicuous; lateral processes conspicuous; antenna with a 3-segmented club; maximal diameter of eye almost as long as or shorter than antennal segment X. Promesonotal dome in dorsal view largely smooth, or sparsely sculptured with weak transverse rugulae, in lateral view with a low to conspicuous mound on its posterior slope; humerus not produced; the dome much narrower at the humeri than at the bottom; mesopleuron, metapleuron and lateral face of propodeum well punctured, sometimes overlain with weak rugulae; propodeal spine narrowly based, usually slightly curved apically. Petiole longer than postpetiole (excluding helcium); postpetiole not massive. First gastral tergite smooth and shining entirely, or sometimes shagreened around its articulation with postpetiole.

Minor — Frons and vertex smooth and shining, or rarely shagreened or very weakly rugoso-punctate; area between antennal insertion and eye weakly rugoso-punctate; preoccipital carina conspicuous dorsally and laterally; median part of clypeus smooth, without a median longitudinal carina; antenna with a 3-segmented club; scape extending far beyond posterolateral margin of head; maximal diameter of eye much shorter than antennal segment X. Promesonotal dome largely smooth and shining, but sometimes weakly rugoso-punctate dorsolaterally, in lateral view with a low or inconspicuous mound on its gentle posterior slope; humerus of the dome in dorsal-oblique view not produced; mesopleuron, metapleuron and lateral face of propodeum well punctured; propodeal spine elongate-triangular, directing upward. Petiole (a little) longer than postpetiole (excluding helcium); postpetiole not massive.

Fischer and Fisher (2013) - Major (n=5): HW 1.13–1.31 (1.25), HL 1.13–1.38 (1.28), SL 0.80– 0.86 (0.83), MDL 0.61–0.71 (0.67), EL 0.16–0.17 (0.17), WL 1.06–1.17 (1.14), PNH 0.41–0.49 (0.45), PNW 0.54–0.59 (0.56), MNH 0.72–0.79 (0.77), PDH 0.34–0.39 (0.36), PTL 0.36–0.40 (0.38), PPL 0.23–0.26 (0.24), PTH 0.22–0.26 (0.24), PPH 0.16–0.22 (0.20), PTW 0.17–0.20 (0.19), PPW 0.31–0.39 (0.34), PSL 0.13–0.17 (0.15), MFL 1.03–1.13 (1.08), MTL 0.77–0.84 (0.82), CI 95–100 (98), SI 61–71 (67), MDI 50–56 (53), EI 13–15 (13), FI 84–91 (87), PSLI 10–14 (12), LPpI 110–144 (120), DPpI 133–150 (143), PpWI 172–195 (183), PpLI 60– 68 (64), PpHI 70–95 (86).

Head almost as wide as long (CI 95–100), sides convex in full-face view. Mandibles smooth, moderately long (MDI 50–56), clypeus smooth, median carina absent and two weak lateral carinae present. Frontal carinae well developed and reaching at least 4/5 of the distance to posterior head margin, antennal scrobe present and punctate. Face longitudinally rugose, interspaces on frons superficially punctate, sides of head and posterolateral lobes with additional cross-meshes between rugae or weakly reticulate, interspaces punctate or weakly punctate. Scapes long (SI 61–71) with decumbent to subdecumbent pilosity and few longer, erect hairs on outer edge. Eyes well developed, moderately large (EI 13–15). Submedian hypostomal teeth and median process small to inconspicuous. Promesonotum high-domed, convex, posteriorly subangulate, in profile with prominent angulate promesonotal process and transverse groove. Dorsum of pronotum mostly smooth with some weak to superficial transverse rugulae, lateropronotum smooth except for weak to superficial curved rugulae, rest of mesosoma punctate, dorsally more weakly so, metanotal groove in profile not impressed or very shallow, with cross-ribs often reduced, dorsum of propodeum in profile usually weakly sloped toward spines and about as long as posterior declivity. Propodeal spines short-spiniform, acute, sometimes very weakly curved posteriorly, slightly shorter than distance between their bases (PSLI 10–14). Metatibia moderately long (FI 84–91), pilosity mostly subdecumbent, along outer edge also with longer suberect hairs. Postpetiole on average 1.4 times wider than long (DPpI 133–150) and 1.8 times wider than petiole (PpWI 172–195), postpetiole shape in dorsal view trapezoidal to hexagonal, sides angulate, ventral process absent or inconspicuous. Dorsum of petiole weakly to superficially punctate, of postpetiole smooth, remainder of both waist segments punctate. Gaster smooth. Standing hairs relatively coarse and abundant, of medium length to moderately long

Minor (n=5): HW 0.52–0.61 (0.58), HL 0.66–0.71 (0.68), SL 0.80– 0.84 (0.81), MDL 0.40–0.44 (0.42), EL 0.13, WL 0.82–0.91 (0.86), PNH 0.30–0.31 (0.30), PNW 0.35–0.40 (0.37), MNH 0.57–0.59 (0.58), PDH 0.26–0.28 (0.27), PTL 0.22–0.30 (0.24), PPL 0.15–0.18 (0.16), PTH 0.14–0.18 (0.16), PPH 0.14–0.16 (0.15), PTW 0.11–0.13 (0.12), PPW 0.19–0.21 (0.19), PSL 0.06–0.08 (0.07), MFL 0.82– 0.90 (0.85), MTL 0.64–0.69 (0.66), CI 79–88 (85), SI 133–154 (140), MDI 70–77 (73), EI 21–25 (22), FI 142–158 (147), PSLI 10–13 (12), LPpI 50–53 (52), DPpI 117–133 (122), PpWI 161–173 (166), PpLI 61–70 (65), PpHI 89– 100 (96).

Head shape roundly oval, longer than wide (CI 79–88), sides convex, posterior head margin relatively wide and weakly rounded, occipital carina visible in full-face view, narrow. Mandibles moderately long (MDI 70–77), longitudinally rugulose. Clypeus smooth, sometimes with short lateral carinae present. Face mostly smooth to superficially punctate, several strong oblique malar carinae present, reaching posterior eye level, sometimes with weak punctures in between, sculpture fading posterior of eyes. Scapes relatively long, about 1.2 times longer than head (SI 133–154), when laid back surpassing posterior head margin by about the length of eleventh funicular segment, with coarse, mostly suberect pilosity. Pronotum in profile flatly convex and subangulate, posterior promesonotal process well developed, bluntly subangulate and prominently produced, metanotal groove well developed and impressed, with weak to inconspicuous cross-ribs, propodeum slightly longer than high, in profile declining smoothly or convexly toward spines. Propodeal spines very short to short-triangular and acute, much shorter than distance between their bases (PSLI 14–19). Promesonotum usually smooth to mostly smooth, remainder of mesosoma punctate to weakly punctate. Metafemur relatively long (FI 142–158), metatibia mostly with subdecumbent to suberect pilosity. Postpetiole usually slightly longer than high (LPpI 100–113), on average 1.7 times wider (PpWI 161–173) and significantly shorter than petiole (PpLI 61–70), with ventral process inconspicuous or very small. Dorsum of petiolar node and postpetiole mostly smooth, remainder weakly to superficially punctate. Gaster smooth and shiny. Standing hairs moderately long, abundant, and blunt, with shorter subdecumbent to suberect pilosity. Color light to darker reddish brown, head usually darker.

Karyotype

  • n = 19, 2n = 38 (Japan) (Imai, 1969) (as Pheidole noda).
  • n = 17 (Crozier, 1975) (as Pheidole noda).
  • n = 18 (Crozier, 1975) (as Pheidole noda).
  • n = 19 (Crozier, 1975) (as Pheidole noda).
  • n = 20 (Crozier, 1975) (as Pheidole noda).

References

References based on Global Ant Biodiversity Informatics

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  • Eguchi K. 2008. A revision of Northern Vietnamese species of the ant genus Pheidole (Insecta: Hymenoptera: Formicidae: Myrmicinae). Zootaxa 1902: 1-118.
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  • Eguchi, K. 2001. A revision of the Bornean species of the ant genus Pheidole (Insecta: Hymenoptera: Formicidae: Myrmicinae). Tropics Monogr. Ser. 2: 1-154.
  • Eguchi, K. 2004. Taxonomic revision of two wide-ranging Asian ants, Pheidole fervens and P. indica (Insecta: Hymenoptera, Formicidae) and related species. Ann. Naturhist. Mus. Wien. 105B:189-209.
  • Eguchi, K. 2008. Eguchi, K. 2008. A revision of North Vietnamese species of the ant genus Pheidole. Zootaxa 1902: 1-118.
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  • Emery, C. "Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei." Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova) (2) 20, no. 40 (1900): 661-722.
  • Emery, C. "Voyage de MM. Bedot et Pictet dans l'Archipel Malais. Formicides de l'Archipel Malais." Revue Suisse de Zoologie 1 (1893): 187-229.
  • Fischer G. & Fisher B. L. 2013. A revision of Pheidole Westwood (Hymenoptera: Formicidae) in the islands of the Southwest Indian Ocean and designation of a neotype for the invasive Pheidole megacephala. Zootaxa 3683: 301-356
  • Fontanilla A. M., A. Nakamura, Z. Xu, M. Cao, R. L. Kitching, Y. Tang, and C. J. Burwell. 2019. Taxonomic and functional ant diversity along tropical, subtropical, and subalpine elevational transects in southwest China. Insects 10, 128; doi:10.3390/insects10050128
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  • Forel A. 1905. Ameisen aus Java. Gesammelt von Prof. Karl Kraepelin 1904. Mitt. Naturhist. Mus. Hambg. 22: 1-26.
  • Forel A. 1906. Les fourmis de l'Himalaya. Bulletin de la Société Vaudoise des Sciences Naturelles 42: 79-94.
  • Forel A. 1912. Ameisen aus Java beobachtet und gesammelt von Edward Jacobson. III. Theil. Notes Leyden Mus. 34: 97-112
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