Ants As Bio Controls

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Ants as biological control agents in agricultural cropping systems
Article in Terrestrial Arthropod Reviews · June 2011

DOI: 10.1163/187498311X571979
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Terrestrial Arthropod Reviews 4 (2011) 157–180 brill.nl/tar
Ants as biological control agents in agricultural

cropping systems
Beth Choate1,2 and Frank Drummond1

1
University of Maine, School of Biology and Ecology, Orono, Maine 04469, USA
2
USDA-ARS, North Central Agricultural Research Laboratory, Brookings, South Dakota 57006, USA
e-mail: Beth.Choate@ars.usda.gov, Frank.Drummond@umit.maine.edu
Received: 16 February 2011; accepted: 14 March 2011
Summary
Ants positively impact agricultural systems by rapidly consuming large numbers of pest insects, disturbing
pests during feeding and oviposition, and increasing soil quality and nutrients. The ability of ants to con-
trol pest species has been recognized since the year 300 A.D. and farmers continue to conserve and pro-
mote ant populations in agricultural systems worldwide. Naturally occurring ant species in milpas, mango,
citrus, coconut, cashews, and cotton control many pest insects. Through judicious insecticide application
and changes in management practices such as tillage, and other manipulations of vegetation and crop
structure, beneficial ant populations are conserved in a variety of agroecosystems. The first recorded exam-
ple of biological control was the manipulation of ants throughout citrus orchards in Asia. Augmentation
continues in citrus, and methods of ant introduction have been developed in Malaysian and Indonesian
cocoa plantations, as well as to control sweet potato and banana weevils in Cuba. Ant species have been
formally incorporated into other integrated pest management programs for cashew in Australia, cocoa in
Papua New Guinea, and mango in Australia and Vietnam. With efforts to reduce chemical pesticide input
in agricultural systems, research evaluating the ability of generalist ant species to control pest insects must
continue.
© Koninklijke Brill NV, Leiden, 2011
Keywords
Ants, Formicidae, biological control, agriculture, integrated pest management, conservation
Introduction
Wilson (1971) estimates that at any given moment there are at least 1015 living ants
moving throughout the earth. Fifteen to twenty percent of the terrestrial animal bio-
mass is comprised of ants; with approximately 12,500 species identified to date
(Hölldobler and Wilson, 1990; Bolton et al., 2006). Due to such great abundance and
total biomass, ants have a major impact on the terrestrial environment. Formicidae
are generalist predators of invertebrate prey (Way and Khoo, 1992), scavengers on
dead animals and feces (Stradling, 1978), fungal farmers (Mueller et al., 1998), they
© Koninklijke Brill NV, Leiden, 2011 DOI 10.1163/187498311X571979
158 B. Choate and F. Drummond / Terrestrial Arthropod Reviews 4 (2011) 157–180
facilitate the aeration of soil and cycling of nutrients (Gotwald, 1986; Hölldobler and
Wilson, 1990), and feed upon and disperse seeds (Rissing, 1981; Beattie, 1985;
Lundgren, 2009). Ants are dominant numerically, geographically, and ecologically,
and their impact on an ecosystem rarely goes unnoticed (Gotwald, 1986). In agroeco-
systems, ants are ideal allies not only as biological control agents, but also as ecosystem
engineers. Below the benefits and potential downfalls of encouraging ants are discussed.
Studies demonstrating their success as predators, methods of conserving and augment-
ing species, and management programs that incorporate ants are presented.
Positive role of ants in agriculture

The ability of ants to rapidly consume large numbers of prey makes them ideal preda-
tors in agroecosystems (Risch and Carroll, 1982). Although colonies may not prey on
all pest insects, disturbance of pests during feeding or oviposition results in reduced
population numbers (Way and Khoo, 1992). In addition to natural pest control, estab-
lishment of ant colonies often leads to improvements in soil quality and increases in
soil nutrients (Gotwald, 1986). Ants may serve as pollinators in systems where tradi-
tional pollinators are absent; however, their role and significance in pollinating agricul-
tural crops remains largely unknown (Gotwald, 1986; Gómez and Zamora, 1992;
Zamora et al., 1996).
Soil modification
Epigaeic ants influence soil movement, nutrient cycling and water availability through-
out their habitats (Lobry de Bruyn, 1999). Through movement of subsoil to the sur-
face, ants increase cation exchange and the amount of available water for plant
absorption (Finnegan, 1974). Movement of soil during nest and tunnel building and
the accumulation of organic matter in the form of ant prey and nest building materials,
results in physical and chemical changes to the soil (Gotwald, 1986; Folgarait, 1998;
Frouz and Jilková, 2008). Physical impacts of ants on soil include an increase in soil
macroporosity and a decrease in bulk density, which results in soil aeration and perme-
ability for water (Frouz and Jilková, 2008). Studies have demonstrated an increase in
organic matter, phosphorous, nitrogen, and potassium in ant nests when compared to
the surrounding soil (Salem and Hole, 1968; Petal, 1978; Mandel and Sorenson, 1982;
Choate, 2010). Ant activity may also neutralize soil pH (Folgarait, 1998; Frouz and
Jilková, 2008), lowering the pH in basic soils and increasing the pH in acidic soils
(Petal, 1980; Frouz and Jilková, 2008). As ants increase the soil organic material, they
create ideal habitats for a variety of biota (Wagner et al., 1997). Pogonomyrmex barbatus
(Smith) (Hymenoptera: Formicidae) nest soil contained significantly greater numbers
of Protozoa, Acari and Collembola than in control soils (Wagner et al., 1997). Similar
results were observed in nests of Messor andrei (Mayr) (Hymenoptera: Formicidae)
with bacteria, fungi, nematodes and microarthropods all more abundant in nests than
in control soil (Boulton et al., 2003). Frouz et al. (2008) report increased soil nutrients
and pH, as well as increased growth rates in Picea abies (L.) Karst (Pinaceae) trees less
B. Choate and F. Drummond / Terrestrial Arthropod Reviews 4 (2011) 157–180 159
than one meter from Formica polyctena Foerster (Hymenoptera: Formicidae) nests.
Wagner (1997) reports an increase in the seed production of Acacia constricta Benth.
(Fabales: Fabaceae) trees closer to Formica perpilosa Wheeler (Hymentoptera:
Formicidae) nests, when compared to those further from nests (Frouz and Jilková,
2008). Other studies suggest that tree growth is negatively impacted by the presence of
ant nests (Frouz and Jilková, 2008). Research evaluating the role of ant nests on pro-
duction in agricultural cropping systems must be conducted in order to fully under-
stand the benefits on plant growth (Frouz and Jilková, 2008). It is likely that ant nest
density is not great enough to impact annual crops and manipulation of the soil is
likely most beneficial in perennial cropping systems.
Pollination
In his book “Insect Pollination of Crops”, Free (1970) identifies six crops that are pos-
sibly pollinated by ants: cashew nut, beet, bird chili, coconut, lychee and cocoa; how-
ever, the pollination ecology of each of these examples is relatively unknown. Philpott
et al. (2006) observed heavier coffee fruits on open branches when compared to ant
exclusion and pollinator exclusion treatments. Studies did not determine whether this
was specifically related to ant pollination or an interaction between ant and flying
insect pollinators (Philpott et al., 2006). Gómez (2000) evaluated the effectiveness of
ants as pollinators of Lobularia maritime (L.) Desv. (Cruciferae). Camponotus micans
(Nylander) (Hymenoptera: Formicidae) accounted for 81.2% of pollinator visits dur-
ing the summer months. Ant-pollinated plants produced more seeds and ant-
pollinated inflorescences produced significantly more flowers, than those pollinated
by winged insects (Gómez, 2000). Hickman (1974) describes the association
between Polygonum cascadense Baker (Polygonaceae) and Formica argentea Wheeler
(Hymenoptera: Formicidae) as a “low-energy system”. Polygonum cascadense is found
on the hot, dry slopes of the Western Cascades of Oregon where F. argentea is required
for cross-pollination and normal seed set. In the field, Hickman (1974) observed that
the flower shape ensures that the ant’s head comes into contact with sticky pollen.
Petersen (1977) reports ant pollination of three plant species in the Colorado alpine
tundra. High wind velocity, cold temperatures and a lack of food resources decrease the
number of flying pollinators, thus high-altitude ants are filling this niche (Petersen,
1977). Faegri and van der Pijl (1971) argue that ants are capable of visiting blossoms
without acquiring pollen. Their small size and hard, smooth body allows them to steal
nectar with no benefit to the plant (Faegri and van der Pijl, 1971). Thus, the role of
ants in pollination remains a mystery and should be a focus of research based upon
their ubiquity in agroecosystems (Gotwald, 1986).
Pest control
Risch and Carroll (1982) list several characteristics that make ants effective and effi-
cient biological control agents. The abundance of ants throughout many agroecosys-
tems makes them ideal in controlling pest insects. Species that form large, stable
populations are capable of reacting quickly to increasing numbers of pests (Way and
Khoo, 1992) through the use of chemical communication and recruitment (Risch and
Carroll, 1982). During periods of decreased primary prey density, ants can survive by
cannibalizing the colony’s brood or feeding on stored prey, making them less likely to
suffer extensive colony losses (Risch and Carroll, 1982). Additionally, ant workers are
thorough and opportunistic foragers, locating pests even in low numbers. As generalist
predators, ants are capable of switching prey species when they have depleted a resource
(Symondson et al., 2002). Ants may also control pest populations through the inter-
ruption of feeding and oviposition (Janzen, 1966; Bentley, 1977). Studies have dem-
onstrated successful manipulation of foragers to increase their contact with pests (Risch
and Carroll, 1982; Huang and Yang, 1987; Campbell, 1990). The first reported exam-
ple of biological control was based upon this concept. Citrus farmers in Asia would
collect predatory ants, introduce them into orchards and provide strips of bamboo to
accommodate movement throughout the trees (Huang and Yang, 1987).
Potential downfalls
Despite having many positive influences throughout agricultural systems, selection of
biological control agents requires thorough evaluation. Presence of ant workers may
result in increased numbers of Hemiptera that damage plants and vector diseases
(Cherrett, 1986; Peng and Christian, 2010). The survival and maintenance of large,
stable ant populations often requires the presence of honeydew-producing hemipter-
ans; however, ant-tending may prevent natural predators and parasitoids from control-
ling populations (Way and Khoo, 1992; McPhee et al., 2008). Research by Campbell
(1990) demonstrated successful augmentation of predatory ant species in jack pine
stands. The increase in ants throughout stands resulted in an increase in hemipteran
numbers, thus damage to trees by hemipterans was greater than that prevented by ant
predators. When fire ants were excluded or suppressed, Hill and Hoy (2003) report
greater control of citrus aphid by parasitoids and Kaplan and Eubanks (2002) report
fewer aphids and increased predation in cotton. Piñol et al. (2009) report conflicting
results, with aphid populations in citrus groves increasing when ants were excluded
from tree branches. While the authors suggest this is most likely the result of other
predators also being excluded (Piñol et al., 2009), it demonstrates the importance of
evaluating the relationship between ants and hemipterans in each agricultural system.
Ant predators may also attack or irritate humans, livestock and other beneficial
organisms (Vinson and Greenberg, 1986; Vandermeer et al., 1990; Peng and Christian,
2010). Solenopsis invicta Buren (Hymenoptera: Formicidae), Solenopsis geminata
(Fabricius) (Hymenoptera: Formicidae), Wasmannia auropunctata (Roger) (Hymenop-
tera: Formicidae), Oecophylla spp., and Anoplolepis gracilipes (Jerdon) (Hymenoptera:
Formicidae) are all efficient predators throughout cropping systems (Brown, 1959;
Sterling, 1978; Sterling et al., 1984; Way and Khoo, 1992); however, all negatively
affect humans. Peng and Christian (2010) report that spraying mango trees with water
prior to harvest decreases Oecophylla sp. activity, suggesting solutions are possible.
Additionally, certain highly predaceous species, such as S. invicta, may be invasive to
the area in which they are serving as efficient biological control agents. The influence
of invasive species on native ant and arthropod communities must be determined prior
to encouragement to fully understand the impact on the ecosystem (Gotelli and Arnett,
2000). Movement of invasive species nests into agricultural systems from surrounding
areas could lead to further destruction of native ant communities. Thus, the use and
management of ants as biological control agents in cropping systems must be carefully
assessed to ensure that the positive impacts outweigh potential negatives.
Pest control by ant populations

Many agricultural systems report the benefits of naturally occurring ant populations
without human intervention to conserve or manipulate populations (Perfecto and
Castiñeiras, 1998) (Table 1). In Nicaragua, numbers of fall armyworm larvae (Spodoptera
frugiperda J.E. Smith (Lepidoptera: Noctuidae)) and corn leafhopper (Dalbulus maidis
De Long & Wolcott (Hemiptera: Cicadellidae)), and damage to maize plants by fall
armyworm were significantly reduced by ant predators in irrigated maize (Perfecto,
1990; Perfecto, 1991). In Mexican milpas, corn, beans and squash are grown in fields
by plowing one-year growth or in forests by cutting and burning vegetation (Risch and
Carroll, 1982). Perfecto and Castiñeiras (1998) state that in areas of milpa production
where pesticides and technical advice are unobtainable, ants often act as the main
source of pest control. In field milpas, the ant community removed 97% of dead
Drosophila flies within 70 minutes of placement, demonstrating the ability of foragers
to quickly respond to prey populations (Risch and Carroll, 1982).
In West Africa, mango farmers that tolerate the presence of native weaver ants,
Oecophylla longinoda (Latreille) (Hymenoptera: Formicidae), throughout trees are
rewarded with significantly better fruit quality (Van Mele, 2008). Weaver ants forage
extensively throughout their habitats, quickly inhabiting new areas and successfully
preying upon or driving away a variety of pest insects (Way and Khoo, 1992). Van
Mele et al. (2007) found that there was a negative correlation between the number
of ant nests and fruit fly (Diptera: Tephritidae) damage. When mangoes were col-
lected from areas with and without ant populations, the number of fruit fly pupae
ranged from 4.0 to 0.67, respectively. Exclusion of ants from trees resulted in an even
greater increase of fruit fly pupae on fruit (with 77.0 pupae on mangoes) (Van Mele
et al., 2007).
Oecophylla smaragdina (Fabricius) (Hymenoptera: Formicidae) is a predatory spe-
cies, native to Northern Australia, managed by coconut growers within the Pacific
Region (Stapley, 1973) and cashew farmers in Australia (Peng et al., 1995). A common
problem in coconut development is immature nutfall, which results from attack by
plant bugs, caterpillars, weevils or rats (Lever, 1969). Trees containing nests of O. sma-
ragdina had no evident nutfall and the spread of this species throughout a coconut
plantation resulted in increased yield (Brown, 1959; Stapley, 1973). In cashew planta-
tions of the Northern Territory of Australia the major pest insects are the tea mosquito
bug, Helopeltis pernicialis Stonedahl et al. (Hemiptera: Miridae), the fruit-spotting
Table 1. Research citing the benefits of ants throughout agricultural cropping systems. (* represent studies
conducted in the USA).
Crop Ant species Pest controlled Highlights References

Cauliflower Pheidole sp. diamondback moth Decreased leaf Agarwal et al., 2007
larvae, black damage by black
cutworm larvae cutworm larvae
Cashew Oecophylla smaragdina tea mosquito bug, Pests controlled, Peng et al., 1995,
fruit-spotting bug, higher quality 1997 a,b
mango tip-borer, nuts produced
leafroller
Citrus Azteca spp. Atta cephalotes Presence prevents Jutsum et al., 1981
defoliation
Cocoa Wasmannia cocoa mirids Noted control Way and Khoo,
auropunctata of pest 1992
Cocoa Anoplolepis longipes Pantorhytes sp., Reduced pest Smith, 1981; Way
Pseudodoniella numbers and Khoo, 1992
laensis
Coconut Wasmannia coconut bug Reduced pest Brown, 1959; Way
auropunctata, numbers and Khoo, 1992
Anoplolepis gracilipes
Cotton Pheidole sp., cotton leaf worm Significantly Perfecto and
Solenopsis sp. reduced pest Castiñeiras, 1998
numbers
*Cotton Solenopsis invicta boll weevil larvae Reduced Sterling, 1978;
pest numbers, Jones and
prevented Sterling, 1979;
economic loss McDaniel and
Sterling, 1979;
Sterling et al.,
1984
*Cowpea, Snap Solenopsis invicta cowpea curculio Reduced numbers Russell, 1980
pea, Lima of pupae in
bean, and Peas the soil
Maize Pheidole radowszkowkii, fall armyworm, Reduced pest Perfecto, 1990;
Solenopsis geminata corn leafhopper numbers, crop Perfecto, 1991
damage
Mango Oecophylla longinoda fruit fly Better fruit quality Van Mele et al.,
2007
Milpas General ant community Drosophila sp. Consumed 97% Risch and
Carroll1982
*Pecan Solenopsis invicta pecan weevil larvae Decreased pest Dutcher and
survival by15% Sheppard, 1981
Rice Solenopsis geminata apple snail Damaged and Yusa, 2001
consumed egg
masses
*Sugarcane Solenopsis invicta sugarcane borer Reduced pest Lofgren, 1986;
numbers, crop Reagan et al.,
damage 1972; Adams
et al., 1981
bug, Amblypelta lutescens Distant (Hemiptera: Coreidae), the mango tip-borer,

Penicillaria jocosatrix Guenee (Lepidoptera: Noctuidae), and the leafroller, Anigraea
ochrobasis Hampson (Lepidoptera: Noctuidae) (Peng et al., 1997a). Peng et al. (1995)
observed the control of all four of these pest species by O. smaragdina. Additionally,
cashew trees with ants produced higher quality nuts than trees without ants. The tea
mosquito bug causes serious damage to trees without insecticides or biological control;
however, trees that contained colonies of O. smaragdina suffered only minor damage
(Peng et al., 1997b).
In 2009, cotton accounted for 14% of worldwide insecticide use, a 5% decline
from that used in the year 2000 (International Cotton Advisory Committee, 2010).
Realizing the potential of natural enemies in this crop is essential to further reduce
insecticide input. Ants are consistently abundant on cotton plants as a result of the
extrafloral nectaries that provide necessary sugar sources (Koptur, 1992; Perfecto and
Castiñeiras, 1998) to foragers. Pheidole spp. in Brazil and Solenopsis spp. in North
America significantly reduce populations of the cotton leafworm in the soil (Perfecto
and Castiñeiras, 1998). Despite its perceived status as one of the worst pests in the
southeastern U.S. (Tschinkel, 2006), where it is invasive, Sterling (1978), Jones and
Sterling (1979), and Sterling et al. (1984) demonstrated that S. invicta reduces boll
weevil damage significantly throughout cotton. Sterling (1978) reports S. invicta con-
sumed up to 85% of boll weevil larvae with a weekly average of 66%. Solenopsis invicta
prevented economic loss due to boll weevil destruction over an 11-year period (Sterling
et al., 1984). Jones and Sterling (1979) introduced red imported fire ants into experi-
mental cotton fields and removed ants from control fields. Fields where ants were
present contained significantly fewer weevil damaged plots than control fields. The
number of emerging weevil adults and the amount of weevil damage did not exceed
17% in June and early July. Weevil damage was over two times more abundant (39%)
in control plots (Jones and Sterling, 1979). McDaniel and Sterling (1982) report a
consumption rate of 0.2 tobacco budworm eggs per individual S. invicta forager with
a total of 711 foragers captured over three sampling dates. McDaniel and Sterling
(1979) obtained similar results collecting 409 red imported fire ants with a mean of
0.4 eggs consumed. More recent research on the influence of red imported fire ants
in cotton demonstrated that this species negatively influences many herbivorous
insects, including pests and beneficials (Eubanks, 2001). Lepidoptera larvae, tarnished
plant bugs and stinkbugs were all suppressed and are major pests throughout cotton
(Eubanks, 2001).
The red imported fire ant has also been recognized as a predator of sugarcane borer,
pecan weevil larvae, and cowpea curculio. Reagan et al. (1972) report sugarcane borer
is responsible for more than 90% of all sugarcane crop losses. Beneficial predators
account for a 25% control of sugarcane borer, and the red imported fire ant is the most
important of these predators (Lofgren, 1986). Louisiana sugarcane fields that were
treated for fire ants had increased infestations of sugarcane borer (53%), as well as
increased damage (69%) (Reagan et al., 1972). Adams et al. (1981) report a slight
increase in sugarcane borer with the elimination of the imported fire ant in Florida.
Survival of pecan weevil larvae varied from 39% under pecan trees where S. invicta
colonies were not present to 24% where ants were abundant (Dutcher and Sheppard,
1981). Cowpea curculio is a common pest of cowpea, snap bean, lima bean and peas
(Capinera, 2009). Russell (1980) reports a significant reduction in pupae at 2.5, 5 and
8 cm soil depth when fire ants were present.
Local cocoa farmers in Cameroon have observed the invasive W. auropunctata con-
trolling cocoa mirids (Way and Khoo, 1992). In the Solomon Islands, both W. auro-
punctata and A. gracilipes decrease populations of the serious pest Amblypelta cocophaga
China (Hemiptera: Coreidae), the coconut bug (Brown, 1959; Way and Khoo, 1992).
In Papua New Guinea, A. gracilipes controls Pantorhytes spp. and Pseudodoniella laensis
Miller (Hemiptera: Miridae), in addition to deterring other ant species that transmit
Phytopthora spp. in cocoa (Smith, 1981; Way and Khoo, 1992).
Additional research identifying ant predators in citrus, rice and cauliflower is located
throughout the literature. In Trinidad citrus orchards, the presence of Azteca spp. in
trees was negatively correlated with the abundance of termites, and positively corre-
lated with scale and coccid pests. Atta cephalotes (Linnaeus) (Hymnoptera: Formicidae)
often defoliate entire citrus trees; however, the presence of Azteca spp. deters the
leafcutter ants from trees. Insecticide application to trees resulted in the death of Azteca
spp. and hemipteran pests, so trees were defoliated by A. cephalotes (Jutsum et al.,
1981). Solenopsis geminata is a reported predator on egg masses of the apple snail, a
major pest that feeds on rice seedlings throughout Asia. Workers damaged and con-
sumed snail egg masses; however, researchers state that ant densities must be increased
to effectively control this pest (Yusa, 2001). Solenopsis geminata is invasive to Asia
(Holway, 2002) and the influence of increased populations on native arthropod com-
munities must be evaluated. Agarwal et al. (2007) observed Pheidole sp. removing
diamondback moth and black cutworm larvae from cauliflower agroecosystems in
India. Ant-exclusion treatments suffered greater leaf damage by black cutworm larvae
than treatments where ants were included (Agarwal et al., 2007). The ability of ants to
naturally reduce pest insects throughout agroecosystems has been reported by farmers,
managers and researchers. Imagine the influence of these populations if encouraged or
manipulated to more efficiently control pests.
Conservation of ant species

The manipulation of land for agriculture often results in a decrease in natural enemies
of arthropod pests as a result of high disturbance levels (Landis et al., 2000). Reductions
in ant biodiversity may be due to changes in microclimate (Hölldobler and Wilson
1990), reduction of food availability due to chemical pesticide use, and species interac-
tions that lead to alterations in community structure (Lobry de Bruyn, 1999).
Conservation biological control in agricultural cropping systems often requires altering
crop management methods (Landis et al., 2000). Reducing insecticide input, manipu-
lating or encouraging diverse vegetation, applying mulch and altering tillage prac-
tices, the amount of shade (Perfecto and Castiñeiras, 1998), and crop structure are all
methods employed by managers to increase ant species diversity. Variations in crop and
land management required for enhancement of ant predators throughout various
ecosystems will be discussed below.
Insecticides
Changes in insecticide use are often a major component of conservation biological
control (Ruberson et al., 1998; Perfecto and Castiñeiras, 1998). In Cuba, sweet potato
farmers have established reservoirs for the non-native, predaceous Pheidole megacephala
(Fabricius) (Hymenoptera: Formicidae) where pesticide applications are prohibited
(Perfecto, 1994). In addition, farmers do not spray synthetic insecticides, but have
adopted the practice of spraying only microbial-based insecticides when necessary
(Perfecto and Castiñeiras, 1998).
Van Mele and Cuc (2000) found that Vietnamese citrus farmers who observed
O. smaragdina in their fields used significantly fewer insecticides, taking advantage of
the ability of this native natural enemy to control pest insects. This species was also
found to be beneficial in Vietnamese sapodilla orchards and its presence resulted in
25% fewer farmers spraying insecticides compared to when this species was absent
(Van Mele and Cuc, 2001). In coconut plantations of the Solomon Islands, selective
application of insecticides resulted in a decrease of the damaging ant species P. mega-
cephala, with no impact on the predaceous, beneficial ant species O. smaragdina
(Brown, 1959; Stapley, 1973).
Altieri and Schmidt (1984) conducted a survey of ant communities in abandoned,
organic and commercially managed apple orchards. Ants were most abundant on the
edge of abandoned fields early in the season, while in conventional fields the number
of ants on the edge increased over time, reaching numbers similar to the organic edges
(Altieri and Schmidt, 1984). Removal of potato tuberworm larvae from baits was
observed to determine ant predation pressure. Ants exerted the greatest predation pres-
sure in abandoned fields, predation decreased by approximately 18% in organic fields,
and by approximately 44% in commercially managed fields. The authors suggest that
management practices such as cultivation and insecticide application are having a
major impact on ant communities throughout this system (Altieri and Schmidt, 1984).
Similar results were observed in Maine lowbush blueberry fields. Ant species richness
was compared between conventionally farmed fields where chemical pesticides were
employed and organically farmed fields. There was a significant increase in both ant
diversity and abundance of the native, highly predaceous Formica exsectoides Forel
(Hymenoptera: Formicidae) in organic fields (Choate, 2010). Laboratory studies
evaluating the impact of low toxicity insecticides on F. exsectoides demonstrated that
microbial and neonicotinoid insecticides may be used in conjunction with this species
to control blueberry pest insects (Choate, 2010). Seagraves and McPherson (2003)
observed variations in the susceptibility of fire ants to common insecticides, with ace-
phate, chlorpyrifos and methomyl being the most toxic to workers. Research evaluat-
ing the effects of insecticides on ant natural enemies is scarce; however, it is a necessary
aspect of the development of integrated pest management programs involving

ant predators.
Vegetational diversity
Perfecto and Castiñeras (1998) list diverse microhabitats, alternative hosts and shelter,
and greater availability of prey, nectar and pollen as factors that contribute to increased
numbers of natural enemies in agricultural cropping systems. In Zanzibar, Way (1954)
observed a greater abundance of predatory O. longinoda colonies in coconut palms
interplanted with other tree species, such as citrus and clove. Interplanting increases
O. longinoda success by providing space for more dense populations, and additional
Hemiptera, as well as providing a more preferred host plant (Way, 1954). Saks and
Carroll (1980) observed a faster removal of baits in a corn, bean, and squash polycul-
ture when compared to a bean monoculture; however, bait removal was fastest in corn
monocultures. The authors suggest this may be a result of the microhabitat created by
vegetation with the tall open canopy of corn providing a very different foraging habitat
than the low, closed canopies of the bean monoculture (Saks and Carroll, 1980). In
Georgia, Tillman et al. (2004) observed greater numbers of invasive, red imported fire
ants in crimson clover interplanted with cotton. In Alabama, Harvey and Eubanks
(2004) report fewer numbers of fire ants in collards intercropped with clover when
compared with collard monocultures; however, predation on diamondback moth lar-
vae was significantly greater in polycultures. In lowbush blueberry fields, Choate
(2010) determined that abundance of F. exsectoides mounds was positively correlated
with the increase of weedy vegetation. It is likely that this mound increase is the result
of greater honeydew-producing Sternorryncha populations on vegetation, which Way
(1954) also suggests contributes to increased Oecophylla sp. nests in coconut.
Perfecto and Sediles (1992) observed no change in ant abundance between maize
monocultures and maize-bean interplantings. Wooline and Reagan (2001) planted
various species of cover crops after sugarcane harvest over a three year period in
Louisiana. No difference in S. invicta abundance was observed between years. During
the final year of the study, abundance of S. invicta workers at baits was significantly
greater in clover plots, than in vetch plots treated with herbicide (Wooline and Reagan,
2001). The role of diverse vegetation in encouraging biological control varies across
cropping systems and must be experimentally manipulated to make management
recommendations.
Tillage
Conventional tillage involves the disturbance of soil-inhabiting arthropods by altering
their distribution, as well as the weed community and the distribution of pesticide resi-
dues (Stinner and House, 1990). A decrease in soil disturbance by reducing or elimi-
nating tillage often increases the presence of beneficial arthropod predators (Stinner
and House, 1990; Perfecto and Castiñeros, 1998). Robertson et al. (1994) found that
when fields were managed under conventional, reduced and zero tillage intensities,
ants were most abundant in fields with zero tillage. This differential in ant abundance
was only observed the final three years of the study (Robertson et al., 1994) most likely
due to the fact that prior to this study all fields had been farmed with conventional
plowing for 25 years.
In fallow fields in Nicaragua, Perfecto (1991) observed that numbers of the native
S. geminata were greater than those of other ant species approximately 20 days after
plowing. Solenopsis geminata remains the dominant species for approximately 30 d,
and then numbers begin to decline. Seven and a half months after plowing, S. geminata
abundance decreased and other ant species abundance increased to numbers observed
immediately after plowing (Perfecto, 1991). Solenopsis geminata can be an important
predator throughout agricultural systems; however, the increase in population num-
bers observed after plowing must coincide with the pest and/or stage of pest being
controlled (Perfecto and Castiñeros, 1998). Increased soil disturbance decreases the
abundance of ants in agroecosystems; however, the effect on individual species may
vary. Managers must determine if increasing diversity or numbers of an individual spe-
cies is most effective in controlling pest populations.
Mulching
In South Carolina, studies evaluating the effects of mulching on the invasive S. invicta
have been conducted. Jackson and Harrison (2008) sampled S. invicta populations in
a killed cover crop conventional tillage system for sweet potato. The four treatments
included killed cover crop left as mulch that was hand-weeded, killed cover crop left as
mulch with no weed control, conventional tillage hand-weeded, and conventional till-
age with no weed control. During the first year of the study, fire ants were most abun-
dant in the killed cover crop treatments than the conventional tillage treatments,
although the difference was not significant. During the second year of the study, fire
ant numbers were significantly increased in the killed cover crop, hand-weeded treat-
ment when compared to both treatments where weedy vegetation was not controlled
(Jackson and Harrison, 2008). Pullaro et al. (2008) observed a significant increase in
fire ant numbers in green bell pepper plots with killed cover crop mulch when com-
pared to plots with black polyethylene mulch alone and black polyethylene mulch with
methyl bromide. The authors suggest this result may be due to the methyl bromide
fumigation or the plastic acting as a barrier for certain ant species to build mounds.
Alternatively, invertebrates were most abundant in the mulched cover crop plots, pro-
viding additional food for fire ants (Pullaro et al., 2008). Similar to vegetational diver-
sity and tillage, mulching provides additional habitat for ant species and according to
the literature reviewed, increases numbers of S. invicta. Further research as to the influ-
ence on species richness is necessary to fully understand the effects of mulching.
Shade
Traditional coffee agroecosystems in Mexico and Central America are cultivated by
replacing the lower strata of the forest with coffee, allowing the overhead tree canopy
to remain (Perfecto and Vandermeer, 1996; Moguel and Toledo, 1999; Philpott et al.,
2008). Much of the current production has transformed these diverse systems to
unshaded monocultures (Perfecto and Vandermeer, 1996; Moguel and Toledo, 1999;
Philpott et al., 2008). Decreases in ant diversity have been observed as a result of altera-
tions in coffee production methods (Nestel and Dickschen, 1990; Perfecto and Snelling,
1995; Perfecto and Vandermeer, 1996; Philpott and Armbrecht, 2006); however, these
studies focus on ground foraging species. Perfecto and Snelling (1995) observed little
to no impact on ant species that forage on coffee bushes. The elimination of shade in
these habitats resulted in greater populations of S. geminata (Perfecto and Vandermeer,
1996). Nestel and Dickschen (1990) demonstrated increased foraging efficiency in
unshaded plots, when compared with shaded plots. Philpott et al. (2008) did not
observe any variation in ant activity level between coffee production systems. In the
same study, removal of ants from coffee plants did not influence overall arthropod
densities, but did result in an increase in Lepidoptera and scale insects (Philpott et al.,
2008). Additionally, this study demonstrated no positive relationship between ants
and herbivory, berry borer damage or crop yields (Philpott et al., 2008). Research by
Vandermeer et al. (2002) and Philpott and Armbrecht (2006) suggest that ants do
protect coffee plants from pest insects. Vandermeer et al. (2002) conducted experi-
ments in a shaded coffee polyculture, placing cabbage butterfly larvae on coffee branches
with ant (Azteca sp.) and no ant treatments. Larvae disappeared much more quickly as
ant density increased on branches, demonstrating the relationship between ant abun-
dance and pest control.
Risch and Carroll (1982) evaluated the effects of shading on ant diversity and forag-
ing in the Mexican milpa system. Solenopsis geminata was much more abundant in field
milpas several days post-planting; however, overall ant species richness was much
greater in forest milpas. The proportion of baits occupied and the removal rates of
Drosophila sp. larvae from these baits were much greater in field milpas than in forest
systems. The reviewed literature demonstrates three different influences of shade on ant
communities. In coffee, a decrease in ground foraging ants and no change in arboreal
ants was reported with a decrease in shade; however, numbers of S. geminata increased.
The tropical fire ant is native to north and central America (Holway, 2002); therefore,
encouragement for pest control may prove beneficial overall.
Crop structure
Peng et al. (1999) experimented with modification of vegetation in order to isolate ant
colonies from one another in Australian cashew. Native O. smaragdina colonies are
extremely territorial with boundary fights being the major factor limiting populations
and their success as biological control agents. Peng et al. (1999) removed overlapping
cashew tree branches to prevent foragers of neighboring colonies from encountering
one another. This simple modification resulted in greater control of major pest insects
(Peng et al., 1999).
Knowledge of ant community dynamics in a cropping system is essential for devel-
oping efficient and sustainable conservation biological control strategies. The influence
of land manipulation and crop structural diversity on individual ant species and diver-
sity varies by species and crop. The effect of these methods on the species to be con-
served must be evaluated.
Augmentation of ant populations

Natural ant abundance in agroecosystems may not be great enough for effective pest
control, thus requiring farmers to introduce additional colonies. The earliest record of
biological control occurred in southern China between 304 A.D. and 877 A.D. (Huang
and Yang, 1987). During this time, native weaver ant nests, O. smaragdina, were col-
lected from natural habitats and sold to citrus farmers to place in their trees. From 985
A.D. to 1401 A.D. hog and sheep bladders were placed near ant nests. The ants were
then collected in these colonized substrates and placed on trees. Since about 1600,
Chinese farmers have been using bamboo strips to create bridges between citrus trees
to encourage foraging throughout orchards (Huang and Yang, 1987). In the 1950’s and
60’s chemicals replaced the role of ants in pest management, but by 1977 the cost of
chemical applications and the abundance of resistant pests prompted farmers to utilize
ants once again. When nests were provided food and shelter, orange yield was similar
to that obtained when insecticides were applied (Huang and Yang, 1987). Today, farm-
ers continue to obtain and establish ant colonies, providing food and shelter for the
ants, and placing bamboo poles between trees to facilitate foraging and protect colo-
nies from competing ant species (Van Mele and Cuc, 2000).
Stapley (1973) describes the introduction of the native (Greenslade, 1972) O. sma-
ragdina into Solomon Island coconut palm plantations where colonies are scarce or
lacking. The soursop tree, Annona muricata (L.) (Annonaceae), is the preferred nesting
site of this species; therefore, Stapley (1973) recommends planting soursop trees
throughout coconut plantations and allowing them to develop for approximately one
year. Once a tree has reached two meters in height and becomes infested with hemi-
pterans, an O. smaragdina nest that is likely to possess a queen is selected and carefully
removed from its host tree. It is then transferred to the soursop in the plantations
where workers immediately begin to form a new nest that remains indefinitely if a
queen is present (Stapley, 1973).
In Malaysia and Indonesia cocoa, the black cocoa ant, Dolichoderus thoracicus
(Smith) (Hymenoptera: Formicidae), reduces damage by mirids; however, abundance
is limited due to a lack of nesting sites. Artificial nests composed of polyethene bags
stuffed with cocoa leaf litter are placed on trees where ants are active. Ants will rapidly
colonize these areas and act aggressively towards adult mirids causing them to fly away.
Thus, the mechanism responsible for reducing mirid damage in cocoa is the result of
disturbance not predation (Chong and Fee, 1989).
In Cuba, farmers have employed the invasive (Holway, 2002) P. megachephala for
the control of sweet potato weevil [Cylas formicarius elegantulus (Summ.) (Coleoptera:
Curculionidae)] and banana weevil [Cosmopolites sordidus (Germ.) (Coleoptera:
Curculionidae)]. Local farmers observed P. megachephala voraciously feeding on sweet
potato weevil larvae (Perfecto, 1994) in natural systems. Ants also interfere with weevil
oviposition on the roots of plants, effectively reducing damage. Fenced areas serve as
reservoirs for this species where insecticide application is prohibited and food
resources in the form of various sugar sources and table scraps are provided (Dlott
et al., 1993; Perfecto and Castiñeiras, 1998). Banana stems or rolled banana leaves act
as temporary nesting sites to transport colonies to sweet potato and banana fields
(Dlott et al., 1993; Perfecto and Castiñeiras, 1998; Lagnaoui et al., 2000). Perfecto and
Castiñeiras (1998) report commercial control of banana and sweet potato weevil
requires at least nine colonies/ha. Lagnaoui et al. (2000) report that a much greater
number of nests (60–110 colonies/ha) are required to maintain sweet potato weevil
populations at low levels (3–5%). It is unclear from the publications as to why recom-
mended numbers vary so greatly; however, it may involve the level of control that
managers are trying to obtain.
In the United States, Campbell (1990) observed native F. exsectoides preying upon
various pests of young pine plantations, such as redheaded pine sawfly, jack pine bud-
worm, gypsy moth, and white pine weevil. In an effort to increase predation by this
species, portable ant colonies were designed and placed throughout jack pine planta-
tions (Campbell et al., 1991). Colonies remained active for three to six weeks when a
queen excluder and 1,500 workers were employed in constructing a colony. Greater
numbers of pest insects were consumed when aphids were present to provide a sugar
source (Campbell, 1990). Hemipterans are a major pest of jack pine also; thus, increas-
ing generalist ant predators in this system did not prove a viable method of pest control
(Campbell, 1990).
Augmentation of ant species for purposes of biological control has proven successful
in a number of tropical agroecosystems. Work by Campbell (1990) demonstrates
one of the potential downfalls of incorporating ants into a cropping system. Diffi-
culties collecting entire colonies and the necessity of acquiring queens and brood
must be considered. Physical and environmental requirements of nests must also
be evaluated prior to introduction. Techniques have proven highly successful in a
variety of agroecosystems; therefore, further research identifying predatory species
that may be easily manipulated, as well as suitable artificial nest materials, should
be conducted.
Integrated pest management programs

Many of the previously described conservation and augmentation efforts could
provide a foundation for the establishment of integrated pest management programs
throughout a variety of crops. In this section, established, active IPM programs
are reviewed (Table 2). Integrated pest management programs that involve predatory
ant species for cocoa, sweet potato, banana, mango, and cashew production are
discussed.
Cocoa
Smith (1981) discusses integrated management for cocoa pests and diseases in Papua
New Guinea recommending that farmers enhance colonies of the crazy ant, an invasive
species, A. gracilipes, to control outbreaks of the cocoa weevil. This ant species also
reduces spreading of Phytophthora disease by outcompeting two common, dominant
Table 2. The locations and cropping systems in which ants are or have been employed as biological
control agents and the specific control strategy.
Location Crop Type of biocontrol Ant species References
Australia cashew conservation Oecophylla smaragdina Peng et al., 1999
IPM Oecophylla sp. Peng and Christian, 2010
Australia mango IPM O. smaragdina Peng and Christian, 2005 a,b
China citrus augmentation O. smaragdina Huang and Yang, 1987;
Van Mele and Cuc, 2000
Cuba sweet conservation Pheidole megacephala Perfecto, 1994; Perfecto
potato and Castiñeiros, 1998
augmentation P. megacephala Perfecto, 1994; Lagnaoui
et al., 2000; Dlott
et al., 2003
IPM P. megacephala, Lagnaoui et al., 2000
Tetramorium guineense
Cuba banana augmentation P. megacephala Perfecto and Castiñeiros,
1998
IPM P. megacephala, Gold et al., 2001
T. guineense
Indonesia cocoa augmentation Dolichoderus thoracicus Chong and Fee, 1989
Malaysia cocoa augmentation D. thoracicus Chong and Fee, 1989
Papua New cocoa IPM Anoplolepis gracilipes Smith, 1981
Guinea coconut conservation O. smaragdina Brown, 1959; Stapley, 1973
augmentation O. smaragdina Stapley, 1973
Vietnam citrus conservation O. smaragdina Van Mele and Cuc, 2000
Vietnam mango IPM O. smaragdina Duc and Hao, 2001
Vietnam sapodilla conservation O. smaragdina Van Mele and Cuc, 2001
ant species that carry infective material from the ground to the canopy (Smith, 1981).
A combination of cultural control, biological control and very limited use of insecticides
are suggested in order to control the variety of pest insects throughout cocoa planta-
tions (Smith, 1981).
Sweet potato
Prior to 1989, Cuban farmers relied heavily on fertilizers, pesticides and petroleum
from the Soviet Union. With the collapse of the Soviet Union and its Eastern European
allies in 1989-1990, input of these products dropped by more than half (Perfecto,
1994), forcing farmers to develop new methods to control the sweet potato weevil.
Obtaining clean planting material, crop rotation, irrigation, sex pheromone traps, host
plant resistance and biological control are all suggested for controlling this pest insect.
Although farmers have been observing the predation of ants on weevils since the 1950’s,
the recent development of portable colonies for two exotic ant species, P. megacephala
and Tetramorium guineense Fabr. (Hymenoptera: Formicidae), has become a major

component of this IPM program (Lagnaoui et al., 2000). The establishment of a reserve
area for this species and the collection of colonies in temporary nests transported to
sweet potato farms have allowed farmers to enhance colonies in their crop (Lagnaoui
et al., 2000).
Banana
Gold et al. (2001) outline various control methods for banana weevil, Cosmopolites
sordidus (Germar) (Coleoptera: Curculionidae) in Africa. This pest has been implicated
as the reason for the disappearance of the East African highland cooking banana from
Uganda and western Tanzania. The IPM program established in Cuba is cited in this
research since P. megacephala and T. guineense are dominant ant species throughout
banana plantations. Laboratory observations of T. guineense and Pheidole spp. remov-
ing banana weevil larvae and eggs from planted corms and pseudostems, respectively,
are the only studies cited which demonstrate the control capabilities of this species,
thus it appears further research is needed to fully incorporate ants into this IPM pro-
gram (Gold et al., 2001).
Mango
Oecophylla smaragdina is a major component in the IPM programs of mango in
Australia and Vietnam (Peng and Christian, 2005a, b; Duc and Hao, 2001). Oecophylla
smaragdina is native to Northern Australia, where it is recognized as the most effective
biological control for the following major pests: mango leafhoppers, red-banded thrips,
fruit spotting bugs, seed weevils, fruit flies, leafrollers, mango tip borers, dimpling
bugs, flower caterpillars and mango planthoppers (Peng and Christian, 2005b).
Additionally, it is credited with effectively controlling minor pests such as case caterpil-
lars, leaf beetles, grasptothus bugs, and tea mosquito bugs. Peng and Christian (2005b)
report that control of leafhoppers and fruit-spotting bugs by O. smaragdina is equal
to or better than when synthetic insecticides are employed. For the dimpling bug
and flower caterpillars, the use of ant predators in conjunction with low toxicity
insecticides was found to be more effective than insecticides alone. Despite the fact
that O. smaragdina have a mutualistic relationship with plant hoppers, parasitoid
abundance on eggs is not impacted; thus the use of ants with low toxicity insecticides
leads to greater control of this species as well (Peng and Christian, 2005b). Yet, there
are problems associated with using this ant species as a biological control, such as their
mutualistic association with scales and mealybugs, formic acid damage to fruit and
annoyance to individuals harvesting fruit. Peng and Christian (2005a, b) describe the
use of low toxicity insecticides to reduce scale and mealybug numbers and not impact
ant populations throughout trees. In addition, the isolation of colonies reduced formic
acid damage to fruit and application of water to trees prior to harvest prompted forag-
ers to go back into their nests (Peng and Christian, 2005b). Duc and Hao (2001)
describe preliminary demonstrations of mango IPM in Vietnam using O. smaragdina.
During demonstrations, colonies were established, but evidence of their success could
not be located.
Cashew
Oecophylla sp. ants are the key element in IPM programs for cashew growers in north-
ern Australia. These programs combine the use of ants with farming tactics and organi-
cally approved pesticides to produce high quality nuts. These programs allow farmers
to produce organic produce and avoid unhealthy chemical applications (Peng and
Christian, 2010).
Integrated pest management programs incorporating predatory ants have been
established throughout a variety of tropical cropping systems. Further research, such as
discussed in sections on conservation and augmentation, must be conducted to incor-
porate ants into temperate agroecosystems. The ubiquity of ants throughout the world
suggests their predatory role is important in a greater number of crops than the five
mentioned here.
Conclusions and future research

The majority of research and IPM programs evaluating or employing ants as biological
control agents have been established in tropical locations, with very few studies identi-
fying potential ant predators in the U.S. (Table 1, Table 2). The role of fire ant preda-
tors was thoroughly researched in cotton in the 70’s and 80’s (Sterling, 1978; Sterling
et al., 1984; Jones and Sterling, 1979; McDaniel and Sterling, 1982) and current
research assessing the relationship between fire ants and various arthropod species has
been conducted (Eubanks, 2001; Wickings and Ruberson, 2011). Single publications
on the influence of fire ants on pecan weevil (Dutcher and Sheppard, 1981) and cow-
pea curculio in Georgia (Russell, 1980), sugarcane borer in Louisiana (Reagan et al.,
1972) and collards in Alabama (Harvey and Eubanks, 2004) were located. More recent
research describing the effects of mulching on predatory ants has been published for
spring bell peppers (Pullaro et al., 2006) and sweet potato (Jackson and Harrison,
2008). Of these few investigations, none identify specific management tactics for
encouraging and harnessing the predatory abilities of ants.
In the U.S., produce standards set by the United States Department of Agriculture
and production contracts with food processing plants often require minimum damage
to produce (Office of Technology Assessment, 1995). Broad-spectrum insecticides are
easy to use and relatively inexpensive throughout the country, thus they are most often
employed to meet these demands (Office of Technology Assessment, 1995). In less
developed countries, farmers may not be able to access or afford synthetic insecticides
and must rely on natural enemy populations to control pests (Ehler, 1998). Whatever
the reason, the over-reliance upon insecticides creates many realized and potential
human health and environmental issues (Office of Technology Assessment, 1995). It is
necessary that scientists conduct research in cropping systems worldwide to iden-
tify potential ant natural enemies and determine their effectiveness in controlling
pest insects.
Integration of predaceous ant species into pest management programs both in the
U.S. and abroad requires that researchers first identify cropping systems with the
potential for manipulation to encourage ant species. Perennial cropping systems seem
to be ideal for incorporating ant natural enemies into management programs due to
their stability. However, as was demonstrated by the cited research, annual systems
managed with conservation tillage and cover crops to enhance ant abundance should
be considered. Within these crops, surveys identifying the ant fauna present must be
conducted. Hölldobler and Wilson (1990) point out that this taxon has greatly been
ignored, with Ward (2010) suggesting that there may be two times more species than
currently identified. Knowledge of the ant fauna, and those species that may prove
most effective as control agents, is essential in harnessing the abilities of these generalist
predators.
Experiments evaluating the ability of a species to control pests include assessments
of prey abundance and crop yields when predator densities are manipulated and
direct observations of predation. Novel techniques, such as molecular gut content
analysis (Harwood et al., 2009), in conjunction with traditional laboratory and field
studies make it possible to tease apart the food web dynamics throughout an agroeco-
system. The ecology of the ant species must also be investigated. Mound phenology,
timing of foraging, as well as interactions with other natural enemies, hemipteran
pests, and humans, influence the ability and feasibility of ant biological control
agents. Colony protein requirements are often determined by the production of
brood, and worker foraging is a direct result of colony needs, thus a complete
understanding of pest control requires knowledge of colony phenology. Lundgren
(2009) discusses the addition of sugar into cropping systems by providing extrafloral
nectar-producing plants and/or food sprays. Nectaries may be introduced by choosing
specific crop varieties, such as cotton (Koptur, 1992; Perfecto and Castiñeiras, 1998),
or planting of cover crops. Incorporation of these concepts into IPM programs with
ant predators may prove positive in reducing the negative association between hemi-
pterans and ants. Ant species that may irritate humans should not be excluded from
consideration. Peng and Christian (2010) describe a fairly simple method to prevent
contact between humans and ants during harvest. Knowledge of ant foraging dynam-
ics, such as time of day and the influence of weather, will make manipulation more
successful.
Methods of conservation or augmentation must be tailored to individual cropping
systems and ant species. Knowledge as to how commonly employed insecticides effect
workers, colony survival and behavior is essential. In lowbush blueberry, low toxicity
insecticides are effective on a variety of pest insects (Yarborough and Drummond,
2010) and laboratory studies demonstrated that these insecticides had little impact on
predatory worker survival (Choate, 2010). Pesticide manufacturing companies have
begun to develop low-toxicity insecticides that may be used in conjunction with ant
natural enemies and it is important to identify their influence, as well as potential for
incorporation in IPM programs with ants. Manipulation of vegetation is also likely to
influence ant species establishment and survival. The maintenance of weedy vegetation
throughout lowbush blueberry fields was found to be essential for F. exsectoides sur-
vival. Blueberry does not contain honeydew-producing pests; therefore, weedy vegeta-
tion was required to support hemipteran communities (Choate, 2010), a necessary
resource for this species (Campbell, 1990).
Augmentation of colonies with portable nests has proved successful in sweet potato,
banana, cocoa, citrus, coconut and jack pine stands. In each of these instances, colonies
were collected from an area and moved to new locations. In order for colonies to estab-
lish and forage, the queen or queens and brood must be collected in addition to work-
ers. Colonies of brood and workers may prove successful for immediate pest control;
however, they are unlikely to provide control beyond the development stage of the
larvae. Mass rearing of ants in the laboratory for use in biological control seems unlikely
with most ant species. Hölldobler and Wilson (1990) discuss the difficulties of estab-
lishing the ideal environmental conditions for copulation to occur. Additionally, nests
require specific environmental conditions that may be difficult to mimic with an artifi-
cial nest. Placement of nests throughout fields to provide appropriate amounts of sun-
light, shade and moisture may also prove challenging. The physical requirements of a
nest must be fully understood to properly augment ant populations with artificial nests.
Further research into the creation of reservoirs for ant colonies (Dlott et al., 1993;
Perfecto and Castiñeras, 1998) and alteration of production practices to encourage and
protect ants seems the most likely avenues for incorporating ants in IPM programs.
Acknowledgements
We would like to thank Andrei Alyokhin, William Glanz, Eleanor Groden, and David
Yarborough for providing comments on an earlier version of this manuscript. Thanks
to the University of Maine Graduate School for providing support through the Maine
Economic Improvement Fund Doctoral Dissertation Fellowship. This is Maine
Agricultural and Forest Experiment Station Publication number 3188.
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Ants as biological control agents in agricultural cropping systems

Article in Terrestrial Arthropod Reviews · June 2011

Beth Choate Francis Drummond

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Ants as biological control agents in agricultural

Beth Choate1,2 and Frank Drummond1

Positive role of ants in agriculture

Pest control by ant populations

Crop Ant species Pest controlled Highlights References

bug, Amblypelta lutescens Distant (Hemiptera: Coreidae), the mango tip-borer,

Conservation of ant species

aspect of the development of integrated pest management programs involving

Augmentation of ant populations

Integrated pest management programs

and Tetramorium guineense Fabr. (Hymenoptera: Formicidae), has become a major

Conclusions and future research

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