Pheidole megacephala

An aggressive species that can form large colonies. Fischer and Fisher (2013): 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.

In Japan, this species occurs on Okinawa Island and southwards in open land, sugar-cane fields and coastal barrens, nesting in the soil, often under stones or logs. Minato et al. (1990) observed hygienic behavior involving the heaping away from nests of bodies of dead colony members by this ant.

Identification
Wilson (2003), in his description of the megacephala species group: The single species of this African complex known from the New World, megacephala, is a tropicopolitan tramp globally spread by human commerce. In the New World, it occurs spottily in disturbed habitats from southern Florida throughout the West Indies to Mexico, Central America, and South America as far as southern Brazil. Its affinities will be unknown until the African Pheidole fauna is better analyzed. If I were handed specimens and told they belonged to a species native to the New World, I would be inclined on the basis of superficial appearance to place megacephala in the tristis group--except for its possession of only 2 hypostomal teeth, unique postpetiole shape, and unicolonial, polygynous colony structure. A good formula for recognizing megacephala at a glance is: heart-shaped head, circular mesonotum, oval postpetiole.

Fischer and Fisher (2013) - 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, short-irregular 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).

Malagasy Region
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 Pheidole decepticon and Pheidole 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 (also see the identification section of these other two species). 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.

Distribution
New World - Widespread although spottily distributed, and sometimes locally very abundant, from southern Florida, Bermuda, and the Bahamas south through the West Indies, southern Mexico, and Central America, to as far south in South America as Santa Catarina, Brazil (Wilson 2003). Detected in southern California in 2014 (see AntWeb) and currently subject of active control measures.

It is widespread in south and central Florida north to St. Augustine (St. Johns Co.), but sporadic, often occurring as a huge polydomous population, for example at the Selby Gardens in Sarasota County. In Hawaii this is the dominant ant in many habitats (Huddleston and Fluker 1968), with disastrous effects on the native arthropods, but in Florida it is a minor species. Pest status: where concentrations of this species occur, large groups of scavenging workers may invade buildings. First published Florida record: Smith 1933. (Deyrup, Davis & Cover, 2000.)

Distribution based on Regional Taxon Lists
Afrotropical Region: Cameroun, Cape Verde, Gambia, Guinea, Ivory Coast, Kenya, Mali, Mozambique, Rwanda, Saudi Arabia, Senegal, Sierra Leone, Uganda, United Arab Emirates, Zimbabwe. Australasian Region: Australia, New Caledonia, New Zealand. Indo-Australian Region: Borneo, Cook Islands, Fiji, Guam, Hawaii, Indonesia, Kiribati, Malaysia, Marshall Islands, Micronesia (Federated States of), New Guinea, Niue, Northern Mariana Islands, Palau, Philippines, Samoa, Solomon Islands, Tokelau, Tonga, Vanuatu, Wallis and Futuna Islands. Malagasy Region: Madagascar, Mauritius, Mayotte, Réunion, Seychelles. Nearctic Region: United States. Neotropical Region: Aruba, Bahamas, Barbados, Bermuda, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, French Guiana, Greater Antilles, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Lesser Antilles, Mexico, Netherlands Antilles, Netherlands Antilles, Nicaragua, Panama, Peru, Puerto Rico, Saint Lucia, Saint Martin (French part), Suriname, Trinidad and Tobago, Venezuela. Oriental Region: India, Nicobar Island, Sri Lanka, Thailand, Vietnam. Palaearctic Region: Canary Islands, China, Egypt, Greece, Iran, Italy, Japan, Montenegro, Oman, Republic of Macedonia, Romania, United Kingdom of Great Britain and Northern Ireland.

Biology
This ant is predaceous and scavenging, and may attend Homoptera for honey dew.

From Wilson (2003): The colonies, which are continuous, with no evident pheromone-based boundaries, and large numbers of fertile queens, are able to reach enormous size. In some areas, especially islands such as Madeira, Culebrita, and the Dry Tortugas, they form a virtually continuous supercolony that excludes most other ant species. They do best in relatively moist, disturbed habitats, thus thrive around human habitations and in cultivated land. Nest sites are highly variable, from within and beneath rotting logs and underneath rocks and sidewalk flagstones to the bark and trunk-based detritus of standing trees. Columns of foragers travel substantial distances from one nest site to another and to food sources. P. megacephala are aggressive toward other species, and war with populations of such locally dominant species as the Indo-Australian weaver ant Oecophylla smaragdina and the cosmopolitan Argentine ant Linepithema humile (= Iridomyrmex humilis). General accounts of this important species are given by Wilson (1971), Holldobler and Wilson (1990), and D. F. Williams et al. (1994). A bibliography of the ant for North America is provided by D. R. Smith (1979). The devastating effect on the native Hawaiian insect fauna was described by the pioneering entomologist R. C. L. Perkins (1913).

While this species nests most commonly outside, when found in buildings they occupy ceilings, wall cavities, power outlets and nearly any suitable space.

Bertelsmeier et al. (2015a, b) examined elements of interspecific aggression, and food resource discovery and dominance, between this species and several other highly invasive ants. In laboratory assays Pheidole megacephala was aggressive when confronted with workers of other invasive ants. Of the group of four species that were found to be aggressive, P. megacephala was the most adept at finding and recruiting to food in a laboratory arena experiment. It was also found to be the least effective aggressor of the group of four invasive ants examined.

Eguchi (2008) - For detailed information on biology and ecological and economic impacts of this species see Reimer et al. (1993), Campbell (1994), Hoffmann (1998), Wetterer (1998), Hoffmann et al. (1999), Vanderwoude et al. (2000), etc.

Western Australia
Though identified from stray specimens (possibly in cargo) much earlier, the pest first made its presence known as a colonist in South Perth in 1942 (May and Heterick 2000). The ant now occupies much of the metropolitan area, and is probably assisting, along with the Argentine Ant (Linepithema humile), with the ongoing demise of Perth’s native ant fauna (Heterick et al. 2002). This pest can also be seen in many cities and towns throughout WA.

Florida
Deyrup, Davis & Cover (2000): Pheidole megacephala, like Linepithema humile, has been in Florida for a long time and is the dominant ant on a few sites. These two species appear to be mutually exclusive, and vie for supremacy in Bermuda (Haskins and Haskins 1965) and Hawaii (Reimer 1994). In Florida, P. megacephala occurs from Tampa through the Florida Keys, with one record from the coast of St. Johns Co. Its rarity in north Florida is probably due to cooler winter temperatures. We have no explanation why it is not a dominant species in tropical and subtropical parts of the state.

Japan
Le Breton et al. (2006) found pupae of this species in an Okinawa Island (southern Japan) population were parasitized by an undescribed Uropodidae mite species. The mite appeared to be a native species in the region hence is believed to have shifted its host choice after the arrival of P. megachepala. Ninety-two percent of the nests sampled (n=75) were infested with mites.

Oman
Sharaf et al (2018) - Oman: A nest series was found under a stone next to a large Rubus sp. tree. Many workers were also found under a pomegranate tree. Another nest series was found under a rock where the soil was dry and loose. Several workers were also occurred in moist soil under a stone next to a date palm tree.

Castes
For additional images see Pheidole megacephala image gallery.

Worker
Minor

Nomenclature

 * edax. Formica edax Forskål, 1775: 84 (w.) EGYPT. Junior synonym of megacephala: Emery, 1892b: 160; Dalla Torre, 1892: 90. [If synonymy correct then edax is the senior name; however, under Art. 23.9 of ICZN (1999) edax is a nomen oblitum.]
 *  megacephala. Formica megacephala Fabricius, 1793: 361 (s.) no locality given. Latreille, 1802c: 232 (q.); Mayr, 1861: 70 (s.w.q.m.); Wheeler, G.C. & Wheeler, J. 1953b: 75 (l.). Combination in Pheidole: Roger, 1863b: 30. Senior synonym of trinodis: Roger, 1863b: 30; of edax: Dalla Torre, 1892: 90; Emery, 1892b: 160 (see note under edax); of perniciosa: Emery, 1915j: 235; of pusilla (and its junior synonyms janus, laevigata Smith, laevigata Mayr): Wheeler, W.M. 1922a: 812; of suspiciosa: Donisthorpe, 1932c: 455; of testacea: Brown, 1981: 530; of agilis: Eguchi, 2008: 56; of bernhardae, gietleni, picata, scabrior: Fischer & Fisher, 2013: 333. [P. megalocephala Schulz, 1906: 155; unjustified emendation.] Current subspecies: nominal plus costauriensis, duplex, ilgii, impressifrons, melancholica, nkomoana, rotundata, speculifrons, talpa. See also: Eguchi, 2001b: 77; Wilson, 2003: 549.
 * trinodis. Myrmica trinodis Losana, 1834: 327, pl. 36, fig. 6 (w.) ITALY. Junior synonym of megacephala: Roger, 1863b: 30.
 * pusilla. Oecophthora pusilla Heer, 1852: 15, pl. 1, figs. 1-4 (s.w.q.m.) PORTUGAL (Madeira I.). Combination in Pheidole: Smith, F. 1858b: 173. Subspecies of megacephala: Emery, 1915j: 235. Senior synonym of janus: Mayr, 1886c: 360; of laevigata Smith: Roger, 1859: 259; Emery, 1915j: 235; of laevigata Mayr: Mayr, 1870b: 981 (footnote). Junior synonym of megacephala: Wheeler, W.M. 1922a: 812.
 * agilis. Myrmica agilis Smith, F. 1857a: 71 (w.) WEST MALAYSIA. Combination in Pheidole: Donisthorpe, 1932c: 449. Junior synonym of megacephala; Eguchi, 2008: 56.
 * janus. Pheidole janus Smith, F. 1858b: 175, pl. 9, figs. 13-17 (s.w.) SRI LANKA. Junior synonym of pusilla: Mayr, 1886c: 360.
 * testacea. Atta testacea Smith, F. 1858b: 168 (s.w.) BRAZIL. Combination in Pheidole: Mayr, 1886c: 360. Junior synonym of megacephala: Brown, 1981: 530.
 * perniciosa. Oecophthora perniciosa Gerstäcker, 1859: 263 (w.) MOZAMBIQUE. [Also described as new by Gerstäcker, 1862: 516.] Combination in Pheidole: Roger, 1863b: 31. Junior synonym of megacephala: Emery, 1915j: 235.
 * suspiciosa. Myrmica suspiciosa Smith, F. 1859a: 148 (w.) INDONESIA (Aru I.). Junior synonym of megacephala: Donisthorpe, 1932c: 455.
 * scabrior. Pheidole megacephala var. scabrior Forel, 1891b: 178 (s.w.) MADAGASCAR. Junior synonym of megacephala: Fischer & Fisher, 2013: 333. See also: Forel, 1897c: 188.
 * picata. Pheidole megacephala var. picata Forel, 1891b: 178 (s.w.) MADAGASCAR. Subspecies of megacephala: Forel, 1895a: 49; of punctulata: Forel, 1897c: 186; Forel, 1905b: 163; Santschi, 1910c: 370. Raised to species: Emery, 1915j: 245; Wheeler, W.M. 1922a: 1019. Junior synonym of megacephala: Fischer & Fisher, 2013: 333.
 * gietleni. Pheidole punctulata r. gietleni Forel, 1905b: 164 (s.w.) MADAGASCAR. Subspecies of picata: Emery, 1915j: 245. Junior synonym of megacephala: Fischer & Fisher, 2013: 333.
 * bernhardae. Pheidole picata var. bernhardae Emery, 1915j: 245 (s.w.) MADAGASCAR. [First available use of Pheidole punctulata r. spinosa var. bernhardae Forel, 1905b: 164; unavailable name.] Junior synonym of megacephala: Fischer & Fisher, 2013: 333.

Type Material


Myrmica agilis

Three worker minor syntypes in. Labelled “35 Malac.” (= Malacca, West Malaysia).

Myrmica suspiciosa

Two worker minor syntypes in. Labelled “Aru.”

Pheidole megacephala

Holotype worker major in. Labelled “Bac. 53.”

Fischer and Fisher (2013) - 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 and the Muséum National d'Histoire Naturelle in Paris. 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).

Taxonomic Notes
Fischer and Fisher (2013) - 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.

Description
Wilson (2003) - Major and minor: in side view, entire postpetiole oval in shape, with all of the ventral margin bulging in a conspicuous convexity, and the node oval, low, and weakly developed; mesonotal convexity absent, the promesonotal profile forming a nearly smooth semicircle; color brownish yellow.

Major: outline of head plus mandibles in full-face view forms a near-perfect heart shape; rugoreticulum present between eye and antennal fossa.

Minor: occiput broad, lacking an occipital collar.

MEASUREMENTS (mm) Major (Grand Bahama Island): HW 1.32, HL 1.32, SL 0.64, EL 0.18, PW 0.60. Minor (Grand Bahama Island): HW 0.54, HL 0.62, SL 0.66, EL 0.12, PW 0.34.

COLOR Major and minor: brownish yellow.

Fischer and Fisher (2013) - Major 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.

(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.

Minor 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.



'''Wilson 2003. Upper: major. Lower: minor. BAHAMAS: Grand Bahama Island. Scale bars = 1 mm.'''

Etymology
Gr L megacephala, large-headed, referring to the major. (Wilson 2003)

Other Resources and Articles
Martindale, S. 2014. Discovery of an aggressive ant variety in Costa Mesa touches off a massive search to see how far they've spread. Orange County Register Identification guide to Invasive Ants of the Pacific Islands
 * [[Media:Baker, A.J., Heraty, J.M.et al. 2019. Inverse dispersal patterns in a group of ant parasitoids.pdf|Baker, A.J., Heraty, J.M., Mottern, J., Hang, J.Z., Hines, H.M., Lemmon, A.R., Lemmon, E.M. 2019. Inverse dispersal patterns in a group of ant parasitoids (Hymenoptera: Eucharitidae: Oraseminae) and their ant hosts. Systematic Entomology 45: 1–19]].
 * Bertelsmeier, C., A. Avril, O. Blight, A. Confais, L. Diez, H. Jourdan, J. Orivel, N. St Germes, and F. Courchamp. 2015a. Different behavioural strategies among seven highly invasive ant species. Biological Invasions. 17:2491-2503. doi:10.1007/s10530-015-0892-5
 * Bertelsmeier, C., A. Avril, O. Blight, H. Jourdan, and F. Courchamp. 2015b. Discovery-dominance trade-off among widespread invasive ant species. Ecology and Evolution. 5:2673-2683. doi:10.1002/ece3.1542
 * Brown, W. L., Jr. 1981. Preliminary contributions toward a revision of the ant genus Pheidole (Hymenoptera: Formicidae). Part I. J. Kans. Entomol. Soc. 54: 523-530 (page 530, senior synonym of testacea)
 * Dalla Torre, K. W. von. 1892. Hymenopterologische Notizen. Wien. Entomol. Ztg. 11: 89-93 (page 90, senior synonym of edax)
 * [[Media:Deyrup, M., Davis, L. & Cover, S. 2000. Exotic ants in Florida.pdf|Deyrup, M., Davis, L. & Cover, S. 2000. Exotic ants in Florida. Transactions of the American Entomological Society 126, 293-325.]]
 * [[Media:do Nascimento, L.E., Amaral, R.R. et al. 2020. Ants as potential mechanical vectors of pathogenic bacteria (10.1093@jmet@jaa062).pdf|do Nascimento, L.E., Amaral, R.R., Ferreira, R.M.dos A., Trindade, D.V.S., do Nascimento, R.E., da Costa, T.S., Souto, R.N.P. 2020. Ants (Hymenoptera: Formicidae) as potential mechanical vectors of pathogenic bacteria in a public hospital in the Eastern Amazon, Brazil. Journal of Medical Entomology 57: 1619–1626.]].
 * Donisthorpe, H. 1932c. On the identity of Smith's types of Formicidae (Hymenoptera) collected by Alfred Russell Wallace in the Malay Archipelago, with descriptions of two new species. Ann. Mag. Nat. Hist. 10(10): 441-476 (page 455, senior synonym of suspiciosa)
 * Eguchi, K. 2001a. A revision of the Bornean species of the ant genus Pheidole (Insecta: Hymenoptera: Formicidae: Myrmicinae). Tropics Monogr. Ser. 2: 1-154. (page 77, see also)
 * Emery, C. 1892c [1891]. Note sinonimiche sulle formiche. Bull. Soc. Entomol. Ital. 23: 159-167 (page 160, senior synonym of edax)
 * Emery, C. 1915i. Les Pheidole du groupe megacephala (Formicidae). Rev. Zool. Afr. (Bruss.) 4: 223-250 (page 235, senior synonym of perniciosa)
 * Fabricius, J. C. 1793. Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tome 2. Hafniae [= Copenhagen]: C. G. Proft, 519 pp. (page 361, soldier described)
 * 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.
 * [[Media:Fournier, D., de Biseau, J.-C.et al. 2016. Social structure and genetic distance (10.1371@journal.pone.0156440).pdf|Fournier, D., de Biseau, J.-C., De Laet, S., Lenoir, A., Passera, L., Aron, S. 2016. Social structure and genetic distance mediate nestmate recognition and aggressiveness in the facultative polygynous ant Pheidole pallidula. PLOS ONE 11, e0156440.]].
 * [[Media:Fujioka, H., Okada, Y. 2019. Liquid exchange via stomodeal trophollaxis (10.1007@s10164-019-00602-9).pdf|Fujioka, H., Okada, Y. 2019. Liquid exchange via stomodeal trophallaxis in the ponerine ant Diacamma sp. from Japan. Journal of Ethology 37, 371–375]].
 * [[Media:Garcia, F.R.M., Ovruski, S.M., Suárez, L., Cancino, J., Liburd, O.E. 2020. Biological control of tephritid fruit flies (10.3390@insects11100662).pdf|Garcia, F.R.M., Ovruski, S.M., Suárez, L., Cancino, J., Liburd, O.E. 2020. Biological control of tephritid fruit flies in the Americas and Hawaii: A review of the use of parasitoids and predators. Insects 11, 662.]].
 * [[Media:Herrera, H.W., Baert, L. et al. 2020. Distribution and habitat preferences of Galápagos ants.pdf|Herrera, H.W., Baert, L., Dekoninck, W., Causton, C.E., Sevilla, C.R., Pozo, P., Hendrickx, F. 2020. Distribution and habitat preferences of Galápagos ants (Hymenoptera: Formicidae). Belgian Journal of Entomology, 93: 1–60.]]
 * [[Media:Heterick 2009b.pdf|Heterick, B. E. 2009. A guide to the ants of South-western Australia. Records of the Western Australian Museum, Supplement 76:1-206.]]
 * Hoffman, B. D., A. N. Andersen, G. J. E. Hill 1999. Impact of an introduced ant on native rain forest invertebrates: Pheidole megacephala in monsoonal Australia. Oecologia 120:595-604.
 * Hölldobler, B. and E. O. Wilson. 1990. The Ants. Cambridge, Mass.: Belknap Press of Harvard U. Press, xii + 732 pp.
 * [[Media:Iwata, K., Eguchi, K. et al. 2005. A case study on urban ant fauna of southern Kyusyu, Japan.pdf|Iwata, K., Eguchi, K., Yamane, S. 2005. A case study on urban ant fauna of southern Kyusyu, Japan, with notes on a new monitoring protocol (Insecta, Hymenoptera, Formicidae). Journal of Asia-Pacific Entomology 8, 263-272.]]
 * [[Media:Katayama, M., Tsuji, K. 2010. Habitat differences and occurrence of native and exotic ants on Okinawa Island (10.1111@j.1479-8298.2010.00400.x).pdf|Katayama, M., Tsuji, K. 2010. Habitat differences and occurrence of native and exotic ants on Okinawa Island. Entomological Science 13, 425–429]].
 * [[Media:Kirschenbaum, R. & Grace, J.K. 2008. Agonistic Responses of the Tramp Ants.pdf|Kirschenbaum, R. & Grace, J.K. 2008. Agonistic Responses of the Tramp Ants Anoplolepis gracilipes, Pheidole megacephala, Linepithema humile, and Wasmannia auropunctata (Hymenoptera: Formicidae). Sociobiology 51, 673-683.]]
 * Latreille, P. A. 1802b. Histoire naturelle générale et particulière des Crustacés et des insectes. Tome 3. Familles naturelles des genres. Paris: F. Dufart, xii + 467 pp. (page 232, queen described)
 * Le Breton, J., G. Takaku, and K. Tsuji. 2006. Brood parasitism by mites (Uropodidae) in an invasive population of the pest-ant Pheidole megacephala. Insectes Sociaux. 53(2):168-171. doi:10.1007/s00040-005-0854-5
 * [[Media:LeBrun, E.G., Plowes, R.M. et al. 2019. Ritualized aggressive behavior reveals distinct social structures (10.1371@journal.pone.0225597).pdf|LeBrun, E.G., Plowes, R.M., Folgarait, P.J., Bollazzi, M., Gilbert, L.E. 2019. Ritualized aggressive behavior reveals distinct social structures in native and introduced range tawny crazy ants. PLOS ONE 14, e0225597]].
 * [[Media:Lee, C.-C., Weng, Y.-M. et al. 2020. Analysis of recent interception records (10.3390@insects11060356 ).pdf|Lee, C.-C., Weng, Y.-M., Lai, L.-C., Suarez, A.V., Wu, W.-J., Lin, C.-C., Yang, C.-C.S. 2020. Analysis of recent interception records reveals frequent transport of arboreal ants and potential predictors for ant invasion in Taiwan. Insects 11, 356]].
 * Mayr, G. 1861. Die europäischen Formiciden. Nach der analytischen Methode bearbeitet. Wien: C. Gerolds Sohn, 80 pp. (page 70, soldier, worker, queen, male described)
 * [[Media:Meurgey, F., Ramage, T. 2020. Challenging the Wallacean shortfall.pdf|Meurgey, F. 2020. Challenging the Wallacean shortfall: A total assessment of insect diversity on Guadeloupe  (French West Indies), a checklist and bibliography. Insecta Mundi 786: 1–183.]]
 * Eguchi, K. 2008. A revision of Northern Vietnamese species of the ant genus Pheidole (Insecta: Hymenoptera: Formicidae: Myrmicinae). Zootaxa. 1902:1-118. [[Media:Eguchi 2008.pdf|PDF]]
 * Perkins, R. C. L. 1913. Introduction. Fauna Hawaiiensis 1(6):i–ccxxvii.
 * Roger, J. 1863b. Verzeichniss der Formiciden-Gattungen und Arten. Berl. Entomol. Z. 7(B Beilage: 1-65 (page 30, senior synonym of trinodis)
 * [[Media:Ruano, F., Tinaut, A. et al. 2000. High surface temeratures select for individual foraging in ants.pdf|Ruano, F., Tinaut, A., Soler, J.J. 2000. High surface temperatures select for individual foraging in ants. Behavioral Ecology 11, 396-404.]]
 * Sarnat, E. M., G. Fischer, B. Guenard, and E. P. Economo. 2015. Introduced Pheidole of the world: taxonomy, biology and distribution. Zookeys. 1-109. doi:10.3897/zookeys.543.6050
 * Sharaf M. R., B. L. Fisher, H. M. Al Dhafer, A. Polaszek and A. S. Aldawood. 2018. Additions to the ant fauna (Hymenoptera: Formicidae) of Oman: an updated list, new records and a description of two new species. Asian Myrmecology. 9:e010004; 1-38. doi:10.20362/am.010004
 * Smith, D. R. 1979. Superfamily Formicoidea. In K. V. Krombein, P. D. Hurd, D. R. Smith, and B. D. Burks, eds., Catalog of Hymenoptera in America North of Mexico, Vol. 2: Apocrita (Aculeata), pp. 1323–1467. Washington, D. C.: Smithsonian Institution Press, pp. i–xvi, 1199–2209.
 * [[Media:Trigos-Peral, G., Abril, S. et al. 2020. Behavioral responses to numerical differences when two invasive ants meet (10.1007@s10530-020-02412-4).pdf|Trigos-Peral, G., Abril, S., Angulo, E. 2020. Behavioral responses to numerical differences when two invasive ants meet: the case of Lasius neglectus and Linepithema humile. Biological Invasions]].
 * Wheeler, G. C.; Wheeler, J. 1953b. The ant larvae of the myrmicine tribe Pheidolini (Hymenoptera, Formicidae). Proc. Entomol. Soc. Wash. 55: 49-84 (page 75, larva described)
 * Wheeler, W. M. 1922j. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bull. Am. Mus. Nat. Hist. 45: 711-1004 (page 812, senior synonym of pusilla (and its junior synonyms janus, laevigata Smith, and laevigata Mayr))
 * Wilson, E. O. 1971. The Insect Societies. Cambridge, Mass.: Belknap Press of Harvard U. Press, x + 548 pp.
 * [[Media:Yu, Y, 2016. Risk of alien species introduction to Ogasawara Islands.pdf|Yu, Y. 2016. Risk of alien species introduction to Ogasawara Islands : Case study of ants at Tokyo Port. World Heritage Studies 1, 86-89.]]

References based on Global Ant Biodiversity Informatics

 * Adenuga A. O. 1975. Mutualistic association between ants and some Homoptera - its significance in cocoa production. Psyche 82: 24-28.
 * Bernard F. 1950. Contribution à l'étude de l'Aïr. Hyménoptères Formicidae. Mém. Inst. Fr. Afr. Noire 10: 284-294.
 * Bernard F. 1953. La réserve naturelle intégrale du Mt Nimba. XI. Hyménoptères Formicidae. Mémoires de l'Institut Français d'Afrique Noire 19: 165-270.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Borowiec L., and S. Salata. 2018. Notes on ants (Hymenoptera: Formicidae) from Gambia (Western Africa). Annals of the Upper Silesian Museum in Bytom Entomology 26: 1-13.
 * Braet Y., and B. Taylor. 2008. Mission entomologique au Parc National de Pongara (Gabon). Bilan des Formicidae (Hymenoptera) recoltes. Bulletin S. R. B. E./K.B.V.E. 144: 157-169.
 * Dejean A., F. Azemar, R. Cereghino, M. Leponce, B. Corbara, J. Orivel, and A. Compin. 2015. The dynamics of ant mosaics in tropical rainforests characterized using the SelfOrganizing Map algorithm. Insect Science doi: 10.1111/1744-7917.12208
 * Diame L., B. Taylor, R. Blatrix, J. F. Vayssieres, J. Y. Rey, I. Grechi, and K. Diarra. 2017. A preliminary checklist of the ant (Hymenoptera, Formicidae) fauna of Senegal. Journal of Insect Biodiversity 5(15): 1-16.
 * Djieto-Lordon, C., A. Dejean, R.A. Ring, B. Aloys. 2005. Ecology of an Improbable Association: The Pseudomyrmecine Plant-ant Tetraponera tessmanni and the Myrmecophytic Liana Vitex thyrsiflora (Lamiaceae) in Cameroon. Biotropica 37(3):421-430
 * 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. 2008. Eguchi, K. 2008. A revision of North Vietnamese species of the ant genus Pheidole. Zootaxa 1902: 1-118.
 * Eidmann H. 1944. Die Ameisenfauna von Fernando Poo. 27. Beitrag zu den Ergebnissen der Westafrika-Expedition. Zool. Jahrb. Abt. Syst. Ökol. Geogr. Tiere 76: 413-490.
 * Emery C. 1915. Les Pheidole du groupe megacephala (Formicidae). Revue Zoologique Africaine (Brussels) 4: 223-250.
 * 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
 * Garcia F.H., Wiesel E. and Fischer G. 2013.The Ants of Kenya (Hymenoptera: Formicidae)Faunal Overview, First Species Checklist, Bibliography, Accounts for All Genera, and Discussion on Taxonomy and Zoogeography. Journal of East African Natural History, 101(2): 127-222
 * IZIKO South Africa Museum Collection
 * Kone M., S. Konate, K. Yeo, P. K. Kouassi, and K. E. Linsenmair. 2012. Changes in ant communities along an age gradient of cocoa cultivation in the Oumé region, central Côte dIvoire. Entomological Science 15: 324339.
 * Kouakou L. M. 2015. Evaluation de la diversite des especes de fourmis anthropophiles, natives, exotiques et potentielles invasives en Cote d'Ivoire. Royal Belgian Institute of Natural Sciences
 * Kouakou L. M. M., K. Yeo, K. Ouattara, W. Dekoninck, T. Delsinne, and S. Konate. 2018. Investigating urban ant community (Hymenoptera: Formicidae) in port cities and in major towns along the border in Côte d’Ivoire: a rapid assessment to detect potential introduced invasive ant species. Journal of Animal and Plant Sciences 36(1): 5793-5811.
 * Kouakou L. M. M., W. Dekoninck, M. Kone, T. Delsinne, K. Yeo, K. Ouattara, and S. Konate. 2018. Diversity and distribution of introduced and potentially invasive ant species from the three main ecoregions of Côte d’Ivoire (West Africa). Belgian Journal of Zoology 148 (1): 83–103.
 * Levieux J., and T. Diomande. 1985. Evolution des peuplements de fourmis terricoles selon l'age de la végétation dans une foret de Cote d'Ivoire intacte ou soumise à l'action humaine. Insectes Sociaux 32(2): 128-139.
 * Lévieux J. 1972. Les fourmis de la savane de Lamto (Côte d'Ivoire): éléments de taxonomie. Bulletin de l'Institut Fondamental d'Afrique Noire. Série A. Sciences Naturelles 34: 611-654.
 * Medler J. T. 1980: Insects of Nigeria - Check list and bibliography. Mem. Amer. Ent. Inst. 30: i-vii, 1-919.
 * Menozzi C. 1939. Hymenoptera Formicidae. Missione Biologica nel Paese dei Borana. 3: 97-110.
 * Menozzi C. 1942. Formiche dell'isola Fernando Poo e del territorio del Rio Muni (Guinea Spagnola). 24. Beitrag zu den wissenschaftlichen Ergebnissen der Forschungsreise H. Eidmann nach Spanisch-Guinea 1939 bis 1940. Zoologischer Anzeiger 140: 164-182.
 * Mikissa J. B., J. H. C. Delabie, J. L. Mercier, and D. Fresnau. 2008. Preliminary Assessment on the Interactions of Wasmannia auropunctata in Native Ant Communities (Hymenoptera: Formicidae) of a Mosaic Gallery Forest/Savannah in Lope National Park, Gabon. Sociobiology 51(1): 207-218.
 * Nousiainen A. 2017. Leaf litter ants in indigenous rainforest and coniferous plantations of the Taita Hills, Kenya. Master Thesis University of Helsinki, 53 pages.
 * Ross S. R. P. J., F. Hita Garcia, G. Fischer, and M. K. Peters. 2018. Selective logging intensity in an East African rain forest predicts reductions in ant diversity. Biotropica 1-11.
 * Ruth M., D. S. Didier, O. J. Marie, B. I. Joseph, A. Akoa, and B. Bilong. 2010. Ants and Phragmanthera capitata (Sprengel), Balle (Loranthaceae) impacts on considerable damages caused on fruit trees of the Ndogbong (Douala, Cameroon) chieftaincys orchard. Journal of Agricultural Extension and Rural Development 2(3): 48-53.
 * Samways M. J. 1990. Species temporal variability: epigaeic ant assemblages and management for abundance and scarcity. Oecologia 84: 482-490.
 * Santschi F. 1910. Formicides nouveaux ou peu connus du Congo français. Annales de la Société Entomologique de France 78: 349-400.
 * Santschi F. 1914. Formicides de l'Afrique occidentale et australe du voyage de Mr. le Professeur F. Silvestri. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 8: 309-385.
 * Santschi F. 1914. Meddelanden från Göteborgs Musei Zoologiska Afdelning. 3. Fourmis du Natal et du Zoulouland récoltées par le Dr. I. Trägårdh. Göteborgs Kungliga Vetenskaps och Vitterhets Samhälles Handlingar. 15: 1-44.
 * Santschi F. 1920. Études sur les maladies et les parasites du cacaoyer et d'autres plantes cultivées à S. Thomé. X. Fourmis de S. Thomé. Extrait des Mémoires publiés par la Société Portugaise des Sciences Naturelles. Lisbonne: Imprimerie de la Librairie Ferin, 4 pp.
 * Santschi F. 1925. Formicidae. Mission Rohan-Chabot. 4(3): 159-168.
 * Santschi F. 1935. Hymenoptera. I. Formicidae. Mission Scientifique de l'Omo 2: 255-277.
 * Santschi F. 1937. Résultats de la Mission scientifique suisse en Angola (2me voyage) 1932-1933. Fourmis angolaises. Revue Suisse de Zoologie. 44: 211-250.
 * Santschi F. 1939. Résultats scientifiques des croisières du navire-école belge, "Mercator". XIV. Formicidae s. lt. Mémoires du Musée Royal d'Histoire Naturelle de Belgique. (2)15: 159-167.
 * Santschi, F. "Résultats de la Mission scientifique suisse en Angola, 1928-1929. Formicides de l'Angola." Revue Suisse de Zoologie 37 (1930): 53-81.
 * Stephens S. S., P. B. Bosu, and M. R. Wager. 2016. Effect of overstory tree species diversity and composition on ground foraging ants (Hymenoptera: Formicidae) in timber plantations in Ghana. International Journal of Biodiversity Science, Ecosystem Services & management 12(1-2): 96-107.
 * Tadu Z., C. Djieto-Lordon, R. Babin, Yede, E. B. Messop-Youbi, and A. Fomena. 2013. Influence of insecticide treatment on ant diversity in tropical agroforestry system: some aspect of the recolonization process. International Journal of Biodiversity and Conservation 5(12): 832-844.
 * Taylor B., N. Agoinon, A. Sinzogan, A. Adandonon, Y. N'Da Kouagou, S. Bello, R. Wargui, F. Anato, I. Ouagoussounon, H. Houngbo, S. Tchibozo, R. Todjhounde, and J. F. Vayssieres. 2018. Records of ants (Hymenoptera: Formicidae) from the Republic of Benin, with particular reference to the mango farm ecosystem. Journal of Insect Biodiversity 8(1): 006–029.
 * Taylor B., and S. F. Adedoyin. 1978. The abundance and interspecific relations of common ant species (Hymenoptera: Formicidae) on cocoa farms in western Nigeria. Bull. Ent. Res. 68: 105-121.
 * Wetterer J. K. 2012. Worldwide spread of the African big-headed ant, Pheidole megacephala (Hymenoptera: Formicidae). Myrmecological News 17: 51-62.
 * Wetterer J. K., X. Espadaler, A. L. Wetterer, D. Aguin-Pombo, and A. M. Franquinho-Aguiar. 2006. Long-term impact of exotic ants on the native ants of Madeira. Ecological Entomology 31: 358-368.
 * Wetterer J. K., X. Espadaler, A. L. Wetterer, D. Aguin-Pombo, and A. M. Franquinho-Aguiar. 2007. Ants (Hymenoptera: Formicidae) of the Madeiran archipelago. Sociobiology 49: 265-297.
 * Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. II. The ants collected by the American Museum Congo Expedition. Bulletin of the American Museum of Natural History 45: 39-269.
 * Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bulletin of the American Museum of Natural History 45: 711-1004
 * Yeo K., T. Delsinne, S. Komate, L. L. Alonso, D. Aidara, and C. Peeters. 2016. Diversity and distribution of ant assemblages above and below ground in a West African forest–savannah mosaic (Lamto, Cote d’Ivoire). Insectes Sociaux DOI 10.1007/s00040-016-0527-6