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Datasheet

Pheidole megacephala (big-headed ant)

Summary

  • Last modified
  • 23 June 2017
  • Datasheet Type(s)
  • Natural Enemy
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Pheidole megacephala
  • Preferred Common Name
  • big-headed ant
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • P. megacephala is highly invasive and can displace native ants, although the degree of invasiveness varies geographically. In Florida, USA it has a sporadic distribution and does not appear to be rapidly spreading (...

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Pictures

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PictureTitleCaptionCopyright
Major worker, lateral view. Costa Rica.
TitleMajor worker
CaptionMajor worker, lateral view. Costa Rica.
CopyrightJohn T. Longino
Major worker, lateral view. Costa Rica.
Major workerMajor worker, lateral view. Costa Rica.John T. Longino
Major worker, facial view. Costa Rica.
TitleMajor worker
CaptionMajor worker, facial view. Costa Rica.
CopyrightJohn T. Longino
Major worker, facial view. Costa Rica.
Major workerMajor worker, facial view. Costa Rica.John T. Longino
Minor worker, lateral view. Costa Rica.
TitleMinor worker
CaptionMinor worker, lateral view. Costa Rica.
CopyrightJohn T. Longino
Minor worker, lateral view. Costa Rica.
Minor workerMinor worker, lateral view. Costa Rica.John T. Longino
Minor worker, facial view. Costa Rica.
TitleMinor worker
CaptionMinor worker, facial view. Costa Rica.
CopyrightJohn T. Longino
Minor worker, facial view. Costa Rica.
Minor workerMinor worker, facial view. Costa Rica.John T. Longino

Identity

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Preferred Scientific Name

  • Pheidole megacephala (Fabricius, 1793)

Preferred Common Name

  • big-headed ant

Other Scientific Names

  • Formica megacephala
  • Pheidole edax Forskål, 1775
  • Pheidole janus Smith F, 1858
  • Pheidole laevigata (Smith, 1855)
  • Pheidole laevigata Mayr, 1862
  • Pheidole perniciosa (Gerstäcker, 1859)
  • Pheidole pusilla (Heer, 1852)
  • Pheidole suspiciosa (Smith F, 1859)
  • Pheidole testacea (Smith F, 1858)
  • Pheidole trinodis (Losana, 1834)

International Common Names

  • English: bigheaded, ant; tramp ant

Local Common Names

  • Australia: coastal brown ant; madeira ant
  • Japan: Tsuya-oozu-ari
  • South Africa: brown house ant

EPPO code

  • PHEIME (Pheidole megacephala)

Summary of Invasiveness

Top of page P. megacephala is highly invasive and can displace native ants, although the degree of invasiveness varies geographically. In Florida, USA it has a sporadic distribution and does not appear to be rapidly spreading (Deyrup et al., 1989; Deyrup, 1991). In contrast, the populations in Australia are invading native habitats and displacing native communities (Reichel and Andersen, 1996; Hoffman et al., 1999; Vanderwoude et al., 2000).

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hymenoptera
  •                         Family: Formicidae
  •                             Genus: Pheidole
  •                                 Species: Pheidole megacephala

Notes on Taxonomy and Nomenclature

Top of page Fabricius originally described P. megacephala as Formica megacephala Fabricius 1793:361. There are a number of subspecies:

- Pheidole megacephala costauriensis Santschi 1915: Ghana

- Pheidole megacephala duplex Santschi 1937: Angola

- Pheidole megacephala ilgi Forel 1907: Ethiopia

- Pheidole megacephala impressifrons Wasmann 1905: South Africa

- Pheidole megacephala melancholica Santschi 1912: Ivory Coast

- Pheidole megacephala nkomoana Forel 1916: Zaire

- Pheidole megacephala rotundata Forel 1894: Mozambique

- Pheidole megacephala scabrior Forel 1891: Madagascar

- Pheidole megacephala speculifrons Stitz 1911: Tanzania

- Pheidole megacephala talpa Gerstäcker 1871: Kenya

Former subspecies of megacephala that are now in the separate species of punctulata include:

- Pheidole punctulata Mayr 1866: South Africa

- Pheidole punctulata atrox Forel 1913: Zaire

- Pheidole punctulata spinosa Forel 1891: Madagascar

- Pheidole punctulata subatrox Santschi 1937: Zaire

The most recent taxonomic treatments of the Pheidole megacephala species group are by Emery (1915, 1919). Disc gel electrophoresis confirmed that Pheidole megacephala rotundata and Pheidole punctulata are distinct species in South Africa (Samways and van Lelyveld, 1982).

Description

Top of page Most species of the genus Pheidole, including P. megacephala, have 12-segmented antennae, the terminal three segments of which are enlarged to form a discrete three-segmented club, and a discrete soldier or major worker caste with an allometrically enlarged head. The most distinctive feature of the species P. megacephala is the grossly swollen postpetiole. Mayr (1861) describes the minor worker, major worker, queen and male, and the larva is described by Wheeler and Wheeler (1953).

Minor worker: head length 0.59 mm, head width 0.53 mm, scape (first long segment of the antenna) length 0.58 mm, Webers length (distance from anterodorsal margin of pronotum to posterior-most extension of metapleural lobes) 0.72 mm (n=1). Head rounded behind; promesonotum evenly arched, mesonotal suture very weak; propodeal spines small but distinct; face and pronotum smooth and shining, katepisternum and lateral propodeum feebly foveolate; postpetiole grossly swollen; dorsal pilosity abundant, long, flexuous, some setae on mesosomal dorsum branched near apex, branches minute; colour brown.

Major worker: head length 1.22 mm, head width 1.21 mm, scape length 0.65 mm (n=1). Face between frontal carinae with parallel longitudinal carinae, space between eyes and frontal carinae punctatorugose overlain with parallel longitudinal carinae, rest of face smooth and shiny; hypostomal margin nearly flat, with pair of very small, low tubercles, located about one-third of the distance from the midline to recessed teeth flanking mandibles (superficially looks like there are no hypostomal teeth); dorsal pilosity abundant; sparse subdecumbent setae projecting from the sides of the head in face view.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

IndonesiaPresentIntroduced Invasive Bolton, 1995
JapanPresentIntroduced Invasive Takahashi et al., 2000
MalaysiaPresentLoke and Lee, 2004
MyanmarPresentIntroduced Invasive Bingham, 1903
PhilippinesPresentIntroducedISSG, 2009
Sri LankaPresentIntroduced Invasive Bolton, 1995

Africa

AngolaPresentNative Invasive Wheeler, 1922
CameroonPresentNative Invasive Wheeler, 1922
CongoPresentNative Invasive Wheeler, 1922
Congo Democratic RepublicPresentNative Invasive Wheeler, 1922
Côte d'IvoirePresent Invasive Babacauh, 1982
EgyptPresent Invasive Bolton, 1995
EritreaPresentNative Invasive Wheeler, 1922
EthiopiaPresentNative Invasive Wheeler, 1922
GabonPresentNativeMikissa et al., 2008
GhanaPresentNative Invasive Wheeler, 1922
GuineaPresentNative Invasive Wheeler, 1922
KenyaPresentNative Invasive Wheeler, 1922
MadagascarPresent Invasive Wheeler, 1922
MalawiPresent Invasive Mercer, 1978
MauritiusPresent Invasive Wheeler, 1922
MozambiquePresentNative Invasive Wheeler, 1922
NigeriaPresentNative Invasive Wheeler, 1922
RéunionPresentIntroducedISSG, 2009
Saint HelenaPresentIntroducedISSG, 2009
Sao Tome and PrincipePresentNative Invasive Wheeler, 1922
SenegalPresentNative Invasive Wheeler, 1922
SeychellesPresent Invasive Wheeler, 1922
Sierra LeonePresentNative Invasive Wheeler, 1922
South AfricaPresentNative Invasive Wheeler, 1922; ISSG, 2009
SudanPresentNative Invasive Wheeler, 1922
TanzaniaPresentNative Invasive Wheeler, 1922
-ZanzibarPresentNative Invasive Wheeler, 1922
UgandaPresentTinzaara, 2005
West AfricaPresent
ZambiaPresent,
ZimbabwePresentNative Invasive Wheeler, 1922

North America

BermudaPresentIntroduced Invasive Kempf, 1972
MexicoPresentIntroduced Invasive Kempf, 1972
USAPresent
-FloridaPresentIntroduced Invasive Smith, 1979
-HawaiiPresentIntroduced Invasive Smith, 1979

Central America and Caribbean

BahamasPresentIntroduced Invasive Kempf, 1972
Costa RicaPresentIntroduced Invasive Kempf, 1972
CubaPresentIntroduced Invasive Kempf, 1972
Dominican RepublicPresentIntroduced Invasive Kempf, 1972
HaitiPresentIntroduced Invasive Kempf, 1972
HondurasPresentIntroduced Invasive Kempf, 1972
JamaicaPresentIntroduced Invasive Kempf, 1972
Puerto RicoPresentIntroduced Invasive Kempf, 1972
Saint Vincent and the GrenadinesPresentIntroduced Invasive Kempf, 1972
Trinidad and TobagoPresentIntroduced Invasive Kempf, 1972

South America

BrazilPresentIntroduced Invasive Kempf, 1972
-AmazonasPresentIntroduced Invasive Kempf, 1972
-BahiaPresentIntroduced Invasive Delabie et al., 1995
-Rio de JaneiroPresentIntroduced Invasive Kempf, 1972
-Sao PauloPresentIntroduced Invasive Kempf, 1972
South Georgia and the South Sandwich IslandsPresentIntroduced Invasive Smith, 1879

Europe

DenmarkPresent
ItalyPresentIntroduced Invasive Bolton, 1995
Portugal
-MadeiraPresentIntroduced19th CWetterer et al., 2007
UKPresent,

Oceania

AustraliaPresentIntroduced Invasive Vanderwoude et al., 2000
-Australian Northern TerritoryPresentIntroduced Invasive Vanderwoude et al., 2000
-New South WalesPresent
-QueenslandPresentIntroduced Invasive Vanderwoude et al., 2000
Cook IslandsPresentIntroduced Invasive Wilson and Taylor, 1967
FijiPresentWard and Beggs, 2007
French PolynesiaPresentIntroduced Invasive Morrison, 1997
GuamPresent
KiribatiPresentIntroducedISSG, 2009
Marshall IslandsPresentIntroducedISSG, 2009
New CaledoniaPresentIntroducedISSG, 2009
New ZealandPresentIntroduced Invasive Berry et al., 1997
NiuePresentIntroducedISSG, 2009
Northern Mariana IslandsPresentIntroducedISSG, 2009
PalauPresentIntroducedISSG, 2009
Papua New GuineaPresentIntroduced Invasive Room, 1975
SamoaPresentIntroduced Invasive Wilson and Taylor, 1967
Solomon IslandsPresentIntroduced Invasive Stapley, 1973
TokelauPresentAbbott et al., 2006
TongaPresent
VanuatuPresentIntroducedISSG, 2009
Wallis and Futuna IslandsPresentIntroducedISSG, 2009

History of Introduction and Spread

Top of page P. megacephala is presumed to be native to Africa because of the extensive geographic variation there (all valid subspecies are from Africa or Madagascar). It appears to have rapidly spread to all parts of the tropics and subtropics beginning in the colonial period and was already widespread in the 1800s. Temporary infestations can occur in the temperate zone when colonies are accidentally introduced to greenhouses and other climate-controlled facilities. For example, the species has been reported from England and Denmark (Collingwood, 1979).

Risk of Introduction

Top of page P. megacephala has a high risk of introduction through nursery stock, where colony fragments can escape detection in soil and plant material. Given its diffuse nesting habits, it can potentially be introduced in almost any transported material in planes, boats or trucks. However, it has already been introduced to just about anywhere that it can be.

Habitat

Top of page P. megacephala generally inhabits disturbed habitats, particularly agricultural and urban areas.

Biology and Ecology

Top of page Physiology and Phenology

P. megacephala is a highly invasive species that is easily transported with human commerce. It is one of several ant species referred to as 'tramp ants', due to their ease of transport and widespread establishment (Passera, 1994). It is unicolonial, meaning it forms large amorphous colonies with multiple nests, multiple fertile queens spread throughout the colony and there are no discrete colony boundaries or colony-specific recognition odours (Wilson, 1971; Hölldobler and Wilson, 1990; Williams, 1994; Wilson, 2003). Populations can fluctuate dramatically (Saunders et al., 1976) and in areas where it is introduced it may gradually replace or be replaced by other invasive ants such as Linepithema humile (Haskins and Haskins, 1965, 1988; Lieberburg et al., 1975).

The feeding habits of this ant are extremely generalized. Laboratory colonies consumed raw fish, various insects, small lizards and especially cooked egg yolk (de la Vega et al., 1984). In a baiting study, fish and lepidopteran larvae were preferred to sucrose or honey solutions (Castiñeiras and Brito, 1983). The workers rapidly distribute lipids, sugars and proteins to the larvae and queens (Sanders et al., 1992). In the field, they are major harvesters of honeydew from Hemiptera and this is a major element of their negative economic impact in agricultural settings. In Cuban citrus groves, the workers build earth tunnels on the trunks and branches, and tend scale insects on the roots and at the branch tips (Castiñeiras and Fernandez, 1983). The major workers usually function in defence, but if the minor workers are depleted, the major workers can execute many of the tasks normally carried out by the minor workers (Wilson, 1984).

Reproductive Biology

As is typical of unicolonial species, most reproduction is probably by the expansion of existing colonies and by long-distance dispersal of fragments of colonies that contain workers, brood and fertile queens. Laboratory observations on P. megacephala show that a queen can revive an experimental nest with only ten newly-emerged minor workers and some pupae. However, a nest with no queen, or with a queen and only major workers, larvae or ten minor workers and no pupae, fails to be revived. This is because the queens and major workers can forage but not feed newly-hatched larvae with liquid food (Chang, 1985). In laboratory colonies, a generation of minor workers takes 34-38 days to complete development at 26-27°C (Chang, 1985).

Environmental Requirements

P. megacephala prefers warm tropical and subtropical environments. It generally inhabits disturbed habitats, particularly agricultural and urban areas, but it is capable of invading mature rainforests in Australia (Heterick, 1997; Hoffman, 1998; Hoffman et al., 1999; Vanderwoude et al., 2000). Although widespread and abundant in lowland areas, they can occur up to an elevation of 2000 m on Hawaiian Islands (Wetterer et al., 1998) but may be restricted to geothermal areas (Wetterer, 1998).

Associations

P. megacephala is associated with a wide variety of honeydew-producing Hemiptera throughout its range. The associations are non-specific but may involve herding, protecting from natural enemies and building protective earth shelters. The effect of this association on host plants is often negative due to the direct withdrawal of plant nutrients, the growth of sooty mould and other fungi on accumulated honeydew, and the transmission of viral plant pathogens. In some cases the effect on host plants can be positive. Bach (1991) found that the presence of ants increased the abundance of coccids on Pluchea indica in Hawaii, but also decreased the amount of honeydew, reducing sooty mould, leaf death and abscission. This resulted in an indirect positive effect on the plant. P. megacephala tends the tettigometrid Hilda patruelis on figs in South Africa. This decreases the parasitism of wasp pollinators and reduces seed predation in the syconium, resulting in a net benefit to the figs (Compton and Robertson, 1988; Cushman et al., 1998).

It is a dominant species in ant mosaics in Africa and Papua New Guinea (Majer, 1972; Leston, 1973; Room, 1975; Majer, 1976; Taylor, 1977). In these circumstances it forms exclusive territories that form a mosaic with other dominant ants.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Bacillus thuringiensis kurstaki Pathogen
Bacillus thuringiensis thuringiensis Pathogen
Beauveria bassiana Pathogen

Notes on Natural Enemies

Top of page In a very general sense, the most important natural enemies of P. megacephala are other dominant ant species. A hymenopteran parasitoid (Eucharitidae, Orasema fraudulenta) has been reared from P. megacephala in the Ethiopian region (Reichensperger, 1913; Wheeler and Wheeler, 1937).

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessions Yes
Land vehicles Yes
Mail Yes
Plants or parts of plants Yes
Soil, sand and gravel Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bark adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Bulbs/Tubers/Corms/Rhizomes adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Fruits (inc. pods) adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Roots adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Seedlings/Micropropagated plants adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Wood adults; eggs; larvae; pupae No Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Flowers/Inflorescences/Cones/Calyx
Leaves
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)

Wood Packaging

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Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
Loose wood packing material No
Non-wood No
Processed or treated wood No
Solid wood packing material with bark No
Solid wood packing material without bark No

Impact

Top of page Negative economic impacts of P. megacephala are primarily due to its interactions with honeydew-producing Hemiptera on crops. Often it is not the direct feeding impact of the Hemiptera that causes economic losses, but rather their transmission of plant viruses and other diseases. P. megacephala increases the abundance of the corn delphacid, Peregrinus maidis, which is a vector of maize viral diseases (Dejean et al., 1997; Dejean et al., 2000). They tend Hilda patruelis on peanuts in Zimbabwe (Weaving, 1980). They tend the sugarcane pest Saccharicoccus sacchari in the Bundaberg area of Queensland, Australia (de Barro, 1990). Honeydew-producing Hemiptera are more abundant on cocoa in Ghana (Campbell, 1994) and Nigeria (Taylor and Adedoyin, 1978) when tended by P. megacephala. The workers spread Phytophthora palmivora (brown pod rot) on cocoa (Evans 1971, 1973; Babacauh, 1982). In Hawaii, USA P. megacephala has a long history of aggravating the effects of Hemiptera on pineapple crops (Illingworth 1917, 1927, 1933, 1935; Beardsley et al., 1982; Reimer et al., 1990). In South Africa, P. megacephala is a widespread pest associated with the increased abundance of Coccoidea on citrus (Samways et al., 1982). In one contradictory study, Jahn and Beardsley (1996) found that under laboratory conditions P. megacephala did not affect populations of pseudococcids on pineapples nor did they move pseudococcids from one plant to another. In general, the workers increase the abundance of Hemiptera by decreasing the effectiveness of the predators and parasitoids (Jahn and Beardsley, 1998; González-Hernández et al., 1999) and interfering with biological control efforts (Cudjoe et al., 1993; Reimer et al., 1993; Jahn and Beardsley, 1994).

P. megacephala can also have a negative impact by displacing beneficial ants. Oecophylla longinoda protects coconut trees from the coconut bug Pseudotheraptus wayi in Tanzania and infestation by P. megacephala can reduce the abundance of Oecophylla (Vanderplank, 1960; Oswald, 1991). A similar situation applies for Oecophylla smaragdina on coconuts in the Solomon Islands (Stapley, 1973). The reduction of P. megacephala with hydramethylnon (Amdro) increases O. longinoda abundance and decreases coconut damage from the coconut bug (Oswald, 1991; Zerhusen and Rashid, 1992) but in one study, higher net revenues occurred when coconut bugs were treated directly with insecticides (Oswald, 1991). Allowing more arboreal interconnections by interplanting citrus makes O. longinoda more resistant to displacement by P. megacephala (Seguni et al., 1999).

Another negative economic impact is the direct damage of agricultural equipment. For example, the workers damage the drip irrigation tubes in sugarcane in Hawaii (Chang et al., 1980).

P. megacephala can have beneficial effects by attacking other insect pests, such as the lilly pilly psyllid (Trioza eugeniae) (Young, 2003), egg masses of the southern green stink bug, Nezara viridula, in Hawaiian macadamia orchards (Jones et al., 2001), eggs of Chilo sacchariphagus, a key pest of sugarcane on Reunion Island (Goebel et al., 1999) and lepidopteran larvae on grain sorghum fields in South Africa (Kfir, 1988).

Detection and Inspection

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The Pacific Invasive Ant Key (PIAKey) manual Pacific Invasive Ants Taxonomy Workshop Manual can both be used in identifying invasive ants in the Pacific region.

Prevention and Control

Top of page Cultural Control and Sanitary Methods

Ploughing can disrupt ant nests and decrease ant abundance in maize fields (Dejean et al., 2000).

Chemical Control

Baits are generally more effective than insecticidal sprays and dusts for controlling ant infestations. In a study of potential bait attractants, P. megacephala showed a significant preference for melezitose over glucose, maltose and trehalose, but not over fructose and sucrose. For oil baits it showed a significant preference for olive oil (Cornelius et al., 1996). Various insecticidal baits have proven to be effective in decreasing ant abundance or eliminating ants, with hydramethylnon (Amdro) usually being the most effective (McEwen et al., 1976; Horwood, 1988; Reimer and Beardsley, 1990; Reimer et al., 1991; Hara and Hata, 1992; Petty and Manicom, 1995). The use of insecticidal baits can decrease ant abundance, but the high application rate required may make control economically prohibitive (Samways, 1985). In some tree crops, sticky barriers can be equally effective and much less expensive (Murray, 1982; Samways, 1985). The bagging of flowers decreased ant infestations in red ginger flowers (Hata et al., 1995).

References

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Abbott KL; Sarty M; Lester PJ, 2006. The ants of Tokelau. New Zealand Journal of Zoology, 33(2):157-164. http://www.rsnz.org/publish/nzjz

Babacauh KD, 1982. Role of insect communities and water in the dissemination of Phytophthora palmivora (Butl.) Butl. emend. Bras. & Griff. in cacao plantations in the Ivory Coast. Cafe Cacao Thé, 26(1):31-36

Bach CE, 1991. Direct and indirect interactions between ants (Pheidole megacephala), scales (Coccus viridis) and plants (Pluchea indica). Oecologia, 87(2):233-239.

Barro PJde, 1990. Natural enemies and other species associated with Saccharicoccus sacchari (Cockerell) (Hemiptera: Pseudococcidae) in the Bundaberg area, southeast Queensland. Journal of the Australian Entomological Society, 29(2):87-88; 6 ref.

Beardsley JW Jr; Su TH; McEwen FL; Gerling D, 1982. Field investigations on the interrelationships of the big-headed ant, the gray pineapple mealybug, and pineapple mealybug wilt disease in Hawaii. Proceedings of the Hawaiian Entomological Society, 24(1):51-67

Berry JA; Green OR; Shattuck SO, 1997. Species of Pheidole Westwood (Hymenoptera: Formicidae) established in New Zealand. New Zealand Journal of Zoology, 24(1):25-33; 17 ref.

Bingham CT, 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. London: Taylor and Francis.

Bolton B, 1995. A New General Catalogue of the Ants of the World. Cambridge, Massachusetts, USA: Harvard University Press.

Campbell CAM, 1994. Homoptera associated with the ants Crematogaster clariventris, Pheidole megacephala and Tetramorium aculeatum (Hymenoptera: Formicidae) on cocoa in Ghana. Bulletin of Entomological Research, 84(3):313-318

Castiñeiras A; Brito A, 1983. Comportamiento de Pheidole megacephala (Hymenoptera: Formicidae: Myrmecinae [sic]) ante deferentes tipos de cebos. Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, 6:63-70.

Castiñeiras A; Caballero S; Rego G; Gonzalez M, 1982. Efectividad tecnico - economico del empleode la hormiga leona Pheidole megacephala en el control del tetuan de boniato Cylas formicarius elegantulus. Ciencia y Tecnica la Agricultura, Protection Plant, 5(Suppl.):103-109.

Castiñeiras A; Calderon A, 1982. Susceptibility of Pheidole megacephala to three microbial insecticides: Dipel, Bitoxobacillin 202 and Beauveria bassiana under laboratory conditions. Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, (Suppl.):61-66.

Castineiras A; Castellanos JA, 1983. Report of Pheidole megacephala (Hymenoptera: Formicidae: Myrmecinae) as a predator of Diatraea saccharalis in Cuba. Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, 6(3):7-10

Castiñeiras A; Fernandez X, 1983. Population increase of Toumeyella cubensis (Homoptera: Coccidae) associated with Pheidole megacephala (Hymenoptera: Formicidae: Myrmecinae). Report. Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, 6(3):11-14

Castiñeiras A; Obregon O; Borges A, 1985. Evaluation of traps for transplanting colonies of Pheidole megacephala (Hymenoptera: Formicidae). Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, 8(2):99-106

Castiñeiras A; Obregon O; Borges A, 1985. Persistence of the biological activity of seven chemical insecticides against Pheidole megacephala, under laboratory conditions. Ciencia y Tecnica en la Agricultura, Proteccion de Plantas, 8(1):29-43

Chang VCS, 1985. Colony revival, and notes on rearing and life history of the big-headed ant. Proceedings of the Hawaiian Entomological Society, 25:53-58

Chang VCS; Ota AK; Sanders D, 1980. Parallel ridge barrier to control ant damage to orifices of drip irrigation tubes. Journal of Economic Entomology, 73(3):403-406

Collingwood CA, 1979. The Formicidae (Hymenoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica, 8:1-174.

Compton SG; Robertson HG, 1988. Complex interactions between mutalisms: ants tending homopterans protect fig seeds and pollinators. Ecology, 69(4):1302-1305

Cornelius ML; Grace JK; Yates JRIII, 1996. Acceptability of different sugars and oils to three tropical ant species (Hymen., Formicidae). Anzeiger für Schädlingskunde Pflanzenschutz Umweltschutz, 69(2):41-43; [17 Ref.].

Cudjoe AR; Neuenschwander P; Copland MJW, 1993. Interference by ants in biological control of the cassava mealybug Phenacoccus manihoti (Hemiptera: Pseudococcidae) in Ghana. Bulletin of Entomological Research, 83(1):15-22.

Cushman JH; Compton SG; Zachariades C; Ware AB; Nefdt RJC; Rashbrook VK, 1998. Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa. Oecologia, 116(3):373-380; 39 ref.

De la Vega R; Diaz G; Palacios ME, 1984. Pheidole megacephala as a predator of Boophilus microplus, qualitative and quantitative aspects. Revista de Salud Animal, 6(4):569-575

Dejean A; Djieto-Lordon C; Durand JL, 1997. Ant mosaic in oil palm plantations of the Southwest Province of Cameroon: impact on leaf miner beetle (Coleoptera: Chrysomelidae). Journal of Economic Entomology, 90:1092-1096.

Dejean A; Orivel J; Durand JL; Ngnegueu PR; Bourgoin T; Gibernau M, 2000. Interference between ant species distribution in different habitats and the density of a maize pest. Sociobiology, 35(1):175-189; 31 ref.

Delabie JHC; Nascimento IC do; Pacheco P; Casimiro AB, 1995. Community structure of house-infesting ants (Hymenoptera: Formicidae) in southern Bahia, Brazil. Florida Entomologist, 78:264-270.

Deyrup M, 1991. Exotic ants of the Florida keys (Hymenoptera: Formicidae). In: Eshbaugh WH, ed. Proceedings of the 4th Symposium on the Natural History of the Bahamas. San Salvador, Bahamas: Bahamian Field Station, 15-22.

Deyrup M; Johnson C; Wheeler GC; Wheeler J, 1989. A preliminary list of the ants of Florida. Florida Entomologist, 72(1):91-101

Dlott J; Perfecto I; Rosset P; Burkham L; Monterrey J; Vandermeer J, 1993. Management of insect pests and weeds. Agriculture and Human Values, 10(3):9-15; 20 ref.

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