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12 Dogs for Monitoring Mealybug Incidence

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14



Semiochemicals in Mealybugs



193



14.13 Future Prospects

• The pheromones for only a few of species

such as Planococcus citri and M. hirsutus

were isolated, identified, and used in other

countries. Use of these pheromones as a monitoring tool will be a great boon for the farmers

to identify the initial stages of infestation.

However, the pheromone for several important invasive species such as Paracoccus marginatus Williams and Granara de Willink and

Phenococcus solenopsis Tinsley needs to be

identified for field monitoring. Besides monitoring the incidence, the pheromone may also

be used to study the spread of the mealybug,

for example, the dispersal pattern of P. marginatus and Pseudococcus jackberdsleyi

Gimpel and Miller can be easily documented

in time and space.

• There are several species of potential invasives such as Phenacoccus manihoti, and the

identification of pheromones for these species

will enable us to monitor the entry of this species into India. It will be worthwhile to isolate,

identify, and synthesize these pheromones for

the quarantine monitoring throughout the

world. Installations at ports, airports, and

other entry points will enable in the early

detection of these mealybugs.

• The technique for the isolation, identification,

and characterization of mealybug pheromones

has been standardized over the years, enabling

us to identify the pheromones for any species.

Moreover, the synthesis of pheromones for a

few species has been accomplished. However,

the ability to synthesize pheromone on a large

scale remains an unfulfilled task resulting in

the use of pheromone only for monitoring,

and not for the mating disruption and mass

trapping. Efforts are needed to develop shorter

synthesis schemes for the effective synthesis

of pheromones in both quality and quantity.

• Often the infestation of complex species of

mealybugs was encountered in many crops,

thus necessitating the identification and use of

generic pheromones. Generic pheromone or



























semiochemicals for the complex pheromone

species in any crop will be advantageous to

the farmers and such generic pheromone or

semiochemical will also be of commercial

success for the entrepreneurs.

For the effective management of mealybug,

mating disruption and inoculative releases of

parasitoids (such as Anagyrus pseudococci)

were considered as effective strategies for the

management of mealybugs in vineyards

(Daane et al. 2008). This will enable environmental friendly, healthy, and safe methods of

mealybug management of the future.

In India, the pheromones were seldom used

though pheromones for several Indian species

have been identified elsewhere. A concerted

effort is needed to use pheromones for

monitoring and for mating disruption of the

mealybug species by the plant protection

experts. The entrepreneurs should take efforts

either to import the pheromone or to develop

facilities for indigenously synthesizing the

pheromones and market at a cheaper rate in

order to guarantee the continuous availability

of pheromones to the farmers.

Wherever the pheromones were not identified

for the species, both indigenous and exotic,

efforts must be made to identify and synthesize

pheromones that can be useful for monitoring

the pests, which can be effective tools for quarantine monitoring and population studies.

Awareness should be brought to the farmers

and the pest management experts on the scope

of using the pheromones for effective management of mealybugs.

Work has to be initiated on the role of plant

volatiles in the attraction of mealybugs and

their natural enemies, which can be used both

for monitoring of pest and natural enemies

and for reinforcing the natural enemy

populations.

Collaborative efforts between countries need

to be made through international funding to

isolate, identify, and synthesize pheromones

of potential invasives for quarantine screening, a prophylactic measure of biosecurity.



194



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of sex pheromone release by the females of two mealybug species (Homoptera Coccoidea Pseudococcidae).

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15



Ant Association

M. Mani and C. Shivaraju



The classic ant–aphid mutualistic relationship

has long been observed by naturalist and entomologist alike, and several studies were conducted on the actual/benefits and factors involved

in these associations. This type of relationship

between ants and other insects is known to occur

in a number of homopterous groups, especially in

the mealybugs. In the case of mealybugs, the

degree of dependence on the ants may vary from

strong and almost necessary associations to

weak, casual seasonal relationships. The association of ants with the mealybugs resulted in the

hypothesis “more the ants, more the mealybugs.”

Ants are often associated with mealybugs as honeydew consumers. Hemiptera-tending ants are

mostly species of the subfamilies Myrmicinae,

Dolichoderinae, and Formicinae (Degen and

Gersani 1989; Mittler and Douglas 2003).

Samways et al. (1982) reported that 11 % of the

123 ant species identified in citrus orchards in

South Africa were associated with mealybugs.

Some mealybugs have an obligatory association

with ants: all Southeast Asian myrmecophilous

mealybugs have been collected only with ants of

the genera Acropyga, Dolichoderus, or

Polyrhachis, which attend the mealybugs either

in subterranean nests or on aerial plant parts

(Gullan and Kosztarab 1997). Aboveground nests

M. Mani (*) • C. Shivaraju

Indian Institute of Horticultural Research,

Bangalore 560089, India

e-mail: mmani1949@yahoo.co.in



were also observed on grapevines in Europe in

association with Phenacoccus aceris (Signoret;

Sforza 2008).



15.1



Benefits to Mealybugs



Benefits derived by mealybugs are more numerous than might be expected.



15.2



Protection from Natural

Enemies



Ants protect mealybugs from their natural enemies; this is a very important and long-realized

aspect of mealybug benefit. Natural enemies are

easily disturbed by movements of the ants. Ants

are naturally hostile to any quick or obviously

harmful movements around the honeydew

sources (Herzig 1938; Nixon 1951). The disruption of the activity of natural enemies by ants provides a temporal refuge for mealybugs (Gutierrez

et al. 2008). Ants have long been known to aggravate mealybug populations and other honeydewproducing insect species by disrupting the natural

biological controls on these species.

Ants deter the natural enemies of mealybugs.

There are numerous examples of ants deterring

the predators and parasites of mealybugs. For

instance, ants also reduce parasitism of the cassava

mealybug,

Phenacoccus

manihoti



© Springer India 2016

M. Mani, C. Shivaraju (eds.), Mealybugs and their Management in Agricultural

and Horticultural crops, DOI 10.1007/978-81-322-2677-2_15



199



M. Mani and C. Shivaraju



200



Matile-Ferrero (Cudjoe et al. 1993). A wide variety of natural enemies are known to prey on pineapple mealybugs. Ants protect mealybugs from

their natural enemies (González-Hernández et al.

1999). In the field, Pheidole megacephala

(Fabricius) had a positive association with

Dysmicoccus neobrevipes Beardsley and a negative association with the predators of mealybugs

(Jahn and Beardsley 2000). Collectively, P. megacephala deters predators from attacking D. neobrevipes. Ants are known to attack the parasitoids

and predators of scales and mealybugs while



Mealybugs and ants on a fruit

of noni (Morinda citrifolia)



attending the sucking pests. In Kenya, C. montrouzieri, released for the control of Planococcus

kenyae (LePelley), was eliminated by ants

(Anderson 1926). The ant Pheidole punctata

(F. Smith) was known to destroy the larvae and

adults of Cryptolaemus montrouzieri Mulsant

preying on Planococcus citri in South Africa

(Kirkpatrick 1927). The ineffectiveness of C.

montrouzieri against Planococcus citri (Risso) at

Liguria (Italy) was due to the attack of ants such

as Tapinoma erratium nigerrimum Oryl and

Iridomyrmex humilis Mayr (Constantino 1935).



Ants attending the mealybugs



The presence of Argentine ant, I. humilis,

appeared to be partly responsible for the failure

of Cryptolaemus to become permanently established in Bermuda (Bennett and Hughes 1959).

In India, the failure of establishment of C. montrouzieri in the mealybug-infested citrus orchards

of Assam was due to the activity of ant Oecophylla

smaragdina (Fab.) (Narayanan 1957). The control of P. citri with Cryptolaemus was made ineffective in the presence of ant attendants (Panis

and Brun 1971). Loss of results with C. montrouzieri was caused by I. humilis in France

(Greathead 1976). The ants Cremaster and

Iridomyrmex were known to prey on C. montrouzieri (Collins and Scott 1982). The failure of

ant control is detrimental to biological control of

citrus mealybug (Singh 1978; Narayanan 1957).

The observation of the protective behavior of P.

megacephala against the attack of C. montrouzieri on the pink hibiscus mealybug



Maconellicoccus hirsutus (Green) showed that

all C. montrouzieri introduced were killed and

removed in 132.5 min. The mealybugs that associated with ants are indeed protected from attack

by their predatory natural enemies, although

mealybugs and ants do not have an intimate association (Lai YiChun and Chang NiannTai 2007).

There was an interaction involving the pink

mealybug Sacchariococcus sacchari, the ant

Camponotus compressus (Fabricius), and the

predator C. montrouzieri in sugarcane (Srikanth

et al. 2001). C. montrouzieri was found more in

numbers and proved successful against the

mealybugs in the absence of ants (Van der Goot

1948; Murray 1982). In South Africa, the control

of ants like I. humilis and Anoplolepis custodiens

F. aided the predator C. montrouzieri to give very

good control of P. citri (Greathead 1971). Poutiers

(1922) also suggested protecting C. montrouzieri

from I. humilis in France.



15



Ant Association



201



Protection of mealybugs from predators



Associations among invasive species of ants

and mealybugs are very important in their success in new locations (Helms and Vinson 2002).

The Argentine ant Linepithema humile (formerly

Iridomyrmex humilis) (Mayr) is an example of an

invasive ant species that is a significant pest in

both natural and managed habitats, and it is commonly associated with mealybug outbreaks

(Daane et al. 2006; Silverman and Brightwell

2008).

Saying that ants “protect” mealybugs from

natural enemies does not necessarily mean that

ants are attacking the natural enemies to save

honeydew as a food resource. Ants deter the natural enemies of mealybugs (Jahn and Beardsley

1994; Rohrbach et al. 1988). The encyrtid parasitoid Anagyrus ananatis Gahan of pineapple

mealybug is not only scared away from when

ants are present but they are also rarely killed by

predators such as the ladybird beetles. When ants

are absent, the parasitoid is highly effective in

lowering the mealybug populations in pineapple

plantings. Ant and mealybug interactions were

studied in a pineapple field near Honolu on the

island of Maui, Hawaii. Big-headed ant Pheidole

megacephala was found to have a positive association with gray pineapple mealybug

Dysmicoccus neobrevipes but no association

with Dysmicoccus brevipes (Cockerell). Sticky

trap collections revealed that D. neobrevipes and

D. brevipes are dispersed by the wind. The positive association between P. megacephala and D.

neobrevipes was not due to ants transporting

mealybugs but could have resulted from ants

deterring natural enemies or removing honeydew

(Gary and Beardsley 2000).



However, some mealybug predators, such as

coccinellids, apparently become tolerated by ants

by mimicking the waxy body cover of the mealybugs (Daane et al. 2007). This condition of parasite–predator adaptation is often observed. The

type of ladybird beetle larva is usually very mealy

in appearance and blends in well with its pseudococcid host. Thus, the parasite or predator species is well adapted to the use of ant-attended

hosts. Ladybird beetle predators are often found

among formicid attendants. Ants seem unable to

recognize the ladybird beetle larva as a predator.

There is, therefore, little doubt that the seemingly

mimetic resemblance of the beetle larva to the

mealybug is an aid to its more perfect predaceous

habitat. Another possible mealybug parasite

adaptation has been noted in a species of fly

larva. This larva apparently remains outside its

host’s body and extracts food externally. This

habit would seem to make it readily susceptible

to the attack by ants. In order to overcome the

problem, the larva is always found completely

hidden beneath the mealybug, and only when the

host insect is removed can the larva be seen.



15.3



Ant Constructions



Ants protect the mealybugs from adverse weather

by building earthen shelters around them and

moving them to protected places; and some ant

species actively construct shelters for mealybugs

that provide some protection from unfavorable

environments and natural enemies (Franco et al.

2000; Helms and Vinson 2002; McLeod et al.

2002). There are two main types of ant construc-



M. Mani and C. Shivaraju



202



tions which are important to mealybugs. The first

is the actual ants’ nest and the second is the socalled carton or ant tent. The latter is not important

in California, although Wheeler (1926) mentioned

several instances of “carton nests” in North

America. Ant-nest isolation of the mealybugs is

considered quite important and has been observed

on several occasions in California. In one instance,

Phenacoccus artemisiae Ehrhorn was found in

some of the upper chambers of a Crematogaster

nest. These mealybug specimens were found at

least 2 ft from the nearest host plant and in two

instances were being transported in the mandibles

of an ant. This particular collection was made in

early February, which is a time of very low insect

activity and high precipitation rate. The mealybugs

were quiescent for the most part and were found in

close contact with one another on the ceiling of ant

nest chambers. It seems likely also that parasite

protection would be important.

Fungus is also a parasite of mealybugs,

although infestation is not normally observed

until the pseudococcid has been mounted; when

examining mealybug preparations, however, fungus infestation is often seen. Ant-nest protection

from a highly humid environment is therefore an

important factor in mealybug welfare and is probably directly connected with protection from fungus contamination. Protection from harsh winter

conditions is perhaps the most important factor of

ant-nest benefit to mealybugs in California.

For the most part, ant tents are important in the

tropical areas, where two primary types are found.

The first is constructed with the silk-forming

glands of the ant larvae. The adult, which has no

silk gland, holds a larva in its mandibles and forces

the immature form to produce its silken product in

the desired area. The ant genus Oecophylla is well

known for this habit. The second type of tent is

made of earth, “paper” formed by the ant, and

leaves. Any or all of these materials may play a

role in the tent construction. The tents are normally built over mealybug colonies which may

have as many as 1,200 individuals. Tent dimensions have been recorded up to 4.5 × 2.3 in.

Mealybug-derived benefits are twofold. First,

the tent provides some protection from direct



drops of rain, although the tent itself is in no way

waterproof. Second, the tent, which has only one

very small entrance, is important in shielding the

mealybugs from large parasites. Apparently, the

mealybug parasites are unable to either find the

tent entrance or push their way through. Although

there are many records of tent parasitism, the rate

of incidence is much lower than where no protection is afforded.



15.4



Removal of Honeydew



Ants prevent the accumulation of honeydew by

consuming it (Jahn and Beardsley 1994; Rohrbach

et al. 1988). Honeydew accumulation, and the

sooty mold that can grow on honeydew, may be

detrimental to mealybugs. These ants, especially

P. megacephala, have been blamed for protecting

them against their natural enemies while removing the excess honeydew produced by the mealybugs (González-Hernández et al. 1999).

One of the direct benefits of ant association to

mealybugs is shown in relation to the production

of honeydew and the subsequent contamination

of the honeydew and source insect with sooty

mold. Because this fungus contaminant is often

the cause of mass destruction of mealybugs, certain adaptations have been made to rid the mealybug of the secretion. Ants remove honeydew

from mealybugs, thereby preventing fungi from

attacking mealybugs, and the removal of honeydew prevents contamination, which may be especially detrimental to first-instar nymphs (Cudjoe

et al. 1993; Daane et al. 2006, 2007; Gullan and

Kosztarab 1997; Moreno et al. 1987). Rohrbach

et al. (1988) hypothesized that honeydew feeding

by ants could benefit mealybugs by preventing

the accumulation of honeydew on the mealybugs

themselves. Presumably, immature mealybugs

get stuck in honeydew and die if ants do not

remove it. Phenacoccus alieni McKenzie is

known to squirt a small globule of honeydew

from the anus to a distance of over 4 in.. This

distance is at least 20 times the total length of the

insect and shows that mealybugs are quite capable of ejecting honeydew to distances well out of



15



Ant Association



the range of their personal contamination. When

ants are in attendance, they remove the honeydew

as described above, thus eliminating the problem

of sooty-mold contamination. Because of this,

ant-attended mealybug colonies are quite often

very dense with little distance between individuals to allow for honeydew ejection.

However, in California’s coastal vineyards,

Argentine ants increased densities of the obscure

mealybug Pseudococcus viburni (Signoret), primarily by removing the honeydew that impedes

the movement of crawlers. Meanwhile, the larvae

of C. montrouzieri successfully forage in patches

of high mealybug density. One hypothesis is that

larvae of C. montrouzieri, being also covered

with waxy structures, successfully mimic mealybugs and avoid detection by ants. Furthermore,

when approached by an ant, the coccinellid larva

stops moving and lowers its body against substrate, thus better resembling a sessile mealybug.

The ants move around the larva, and stroke it

with their antennae like they stroke the mealybug. After failing to obtain the honeydew, the ant

moves away. Densities of C. montrouzieri were

higher on ant-tended vines, where there were

more mealybugs (Daane et al. 2007).

Ants stimulated increased feeding by mealybugs; tending by the ants may have other effects

that alter mealybug densities: It may also improve

the mealybugs’ habitat or fitness (Daane et al.

2007). In the presence of ants, mealybugs are

able to ingest larger quantities of sap (Degen and

Gersani 1989). In several instances, ant-tended

mealybug colonies will be much larger than colonies of the same mealybug species on the same

host that are not tended by ants. Therefore, outwardly the ant’s presence must be of some benefit

to the mealybug, either directly or indirectly.



15.5



Transportation of Mealybugs



Ants are known to transport the mealybugs from

plant to plant between and within fields, thus

facilitating mealybug dispersal. In California, it

is often possible to see ant Camponotus actually

carrying from its host plant, directly into the ants



203



nest. Ants are the primary or sole means of

mealybug dispersal in pineapple. Illingworth

(1931) observed P. megacephala carrying mealybugs from one cage of pineapples to another. The

big-headed ant Pheidole megacephala. (F.),

Argentine ant Linepithema humile (Mayr), and

fire ant Solenopsis geminata (F.) are commonly

found in the Hawaiian pineapple agroecosystem,

where they tend pink pineapple mealybugs

(PPM) and gray pineapple mealybugs (GPM) for

honeydew. These ants, especially P. megacephala, have been blamed for dispersing mealybugs

(González-Hernández et al. 1999).



15.6



Benefit to Ants



Access to honeydew has been shown to enhance

the rate of increase of ant colonies. Honeydew

accounts for more than half of the diet of many

temperate wood ants (Formica spp.), and it is the

dominant food source of some subterranean ants

(Mittler and Douglas 2003). Mealybug exudates/

honeydew is highly acidic (pH 3) with fructose

(45 g), glucose (20 %), and other sugar contents

in negligible quantity (0–2 %) per 100 g of solids

(Ashbolt and Inkerman 1990). The normal ant–

mealybug association when observed in the field

is seemingly quite simple. The ants, which may

be of various genera, normally move busily from

one mealybug specimen to another. When a

mealybug is contacted, the ant begins to fondle

the mealybug with its antennae just as it might

fondle its own brood.

The ants rest their heads on the dorsum of the

mealybugs near the area of the ostioles for honeydew. Mealybugs extrude a solution from their

ostioles when disturbed. This is possibly a

defense mechanism. When Phenacoccus echeveria McKenzie is purposefully disturbed, it

extrudes two small globules of honeydew through

its posterior ostioles. An ant then comes along

and within a few seconds ingests all of the

extruded honeydew. It is clear that ants benefit

from mealybugs in receiving honeydew from

them, which is added to the ants’ food supply.

The amount that ants rely on honeydew for their



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