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M. Mani, C. Shivaraju (Eds.) - Mealybugs and Their Management in Agricultural and Horticultural Crops - Springer India (2016)
M. Mani, C. Shivaraju (Eds.) - Mealybugs and Their Management in Agricultural and Horticultural Crops - Springer India (2016)
Shivaraju Editors
Mealybugs
and their
Management in
Agricultural and
Horticultural crops
Mealybugs and their Management
in Agricultural and Horticultural crops
M. Mani C. Shivaraju
Editors
Springer (India) Pvt. Ltd. Springer is part of Springer Science+Business Media (www.springer.
com)
Dedicated to my wife Vijayarani who was
involved with the work on mealybugs.
She had helped me to carryout extensive
surveys for mealybugs in different crops in
India. She has also played a major role in
nalising the draft on Mealybugs and their
Management in Agricultural and
Horticultural crops.
Foreword
vii
Preface
ix
Acknowledgements
xi
Contents
1 Introduction ................................................................................... 1
M. Mani and C. Shivaraju
Part I Mealybugs
2 Morphology ................................................................................... 7
M. Mani and C. Shivaraju
3 Cytogenetics ................................................................................... 19
Ramakrishna Sompalaym, Kokilamani A. Lingarajaiah,
Raju G. Narayanappa, Jayaprakash,
and Venkatachalaiah Govindaiah
4 Taxonomy ....................................................................................... 55
M. Mani
5 Molecular Identification of Mealybugs ....................................... 75
K.B. Rebijith, R. Asokan, and N.K. Krishna Kumar
6 Biology............................................................................................ 87
M. Mani and C. Shivaraju
7 Culturing of Mealybugs ................................................................ 107
M. Mani and C. Shivaraju
8 Mode of Spread of Mealybugs ..................................................... 113
M. Mani and C. Shivaraju
9 Damage........................................................................................... 117
M. Mani and C. Shivaraju
10 Mealybugs as Vectors .................................................................... 123
R. Selvarajan, V. Balasubramanian, and B. Padmanaban
11 Economic Importance................................................................... 131
M. Mani and C. Shivaraju
12 Ecology ........................................................................................... 141
M. Mani and C. Shivaraju
13 Natural Enemies of Mealybugs .................................................... 149
A.N. Shylesha and M. Mani
xiii
xiv Contents
xix
xx List of Tables
xxiii
Introduction
1
M. Mani and C. Shivaraju
Mealybugs belong to the insect group that is fungus called sooty due to which a signicant
commonly known as scale insects; They have infestation of mealybug creates a black, sticky
soft segmented oval bodies, but without an outer mess. Most of the economically important mealy-
shell. Mealybugs (Hemiptera, Sternorrhyncha, bug species are known to be associated with
Coccoidea, Pseudococcidae, and Putoidae) are long lists of host plants, and the development of
small, soft-bodied plant sap-sucking insects. The high population density, which eventually would
name mealybug is descriptive of the insects kill the host plant. Plant growth conditions may
body, which is covered by a white sticky powder strongly affect the development of the mealybug.
resembling cornmeal. Their common name is Flowering and fruiting phases of plant support
derived from the mealy wax secretion that usu- heavy mealybug population. Likewise, hot
ally covers their bodies (Kosztarab and Kozr weather favors rapid multiplication resulting in
1988). Because of their appearance, mealybugs the outbreak of mealybug population.
are often confused for cushionscale insects or Many of the mealybugs show sexual dimor-
woolyaphids. Unlike their close relative scale phism but parthenogenetic mode of reproduction
insects, mealybugs retain their legs throughout is also observed in some species of mealybugs.
their lives. Mealybugs may be oviparous or viviparous or
Mealybugs feed on a variety of herbaceous ovoviparous. The eggs are usually laid in loose
and woody plants, including the angiosperm, masses of cottony wax or felt-like ovisacs. Some
gymnosperm, and fern families. Most of the species bear living young. Only newly hatched
mealybugs are arboreal and some are subterra- mealybugs, also called as crawlers, are not cov-
nean feeding on the roots. They are phloem feed- ered with wax coating, moving from one part to
ers and suck the sap from all plant parts and also another within the plant and also between plants;
transmit some plant disease, thus causing serious this is the most vulnerable stage for chemical
economic losses to economically important crop control. They are windblown, and the spreading
plants. Mealybugs take in great quantities of plant of mealybugs is facilitated by wind. Within two
uids and therefore excrete a lot of liquid waste days, they are also covered by waxy coating,
called honey that supports the growth of a black making them hard to get killed with chemicals.
There are three nymphal instars in female and
four in male mealybugs also covered with wax.
M. Mani (*) C. Shivaraju
Adult male and female mealybugs are completely
Indian Institute of Horticultural Research,
Bangalore 560089, India different from each other. Adult female mealy-
e-mail: mmani1949@yahoo.co.in bugs are characteristically elongate, oval, soft,
Only very few books are available on Ben-Dov Y (2006) Scales in a family/genus query.
Family pseudococcidae & genus. Available at http://
mealybugs: Mealybugs of California by
www.sel.barc.usda.gov/calecgi/chklist.exe? Accessed
McKenzie (1967), Australian Mealybugs by 14 Aug 2008
Williams (1985), Mealybugs of Central and Charles JG, Cohen D, Walker JTS, Forgie SA, Bell VA,
South America by Williams and Granara de Breen KC (2006) A review of grapevine leafroll asso-
ciated virus type 3 (GLRaV-3) for the New Zealand
Willink (1992), A Systematic Catalogue of the
wine industry. The Horticulture and Food Research
Mealybugs of the World (Insecta, Homoptera, Institute of New Zealand Ltd, Auckland
Coccoidea, Pseudococcidae and Putoidae): With Daane KM, Almeida RPP, Bell VA, Walker JTS,
Data on Geographical Distribution, Host Plants, Botton M, Fallahzadesh M, Mani M, Miano JL,
Sforza R, Walton VM, Zaveizo T (2012) Biology
Biology and Economic Importance by Ben-Dov
and management of mealybugs in vineyards. In:
(1994), and Mealybugs of Southern Asia by Bostman NJ et al (ed) Arthropod management in
Williams (2004). They deal mostly with the taxo- vineyard pests, approaches, and future directions.
nomical aspects of mealybugs in different Springer Science + Media B.V., pp 271307. doi:
10.1007/978-007-4032-7-12
regions. Efforts have been made to present infor-
Downie DA, Gullan PJ (2005) Phylogenetic congruence
mation comprehensively about all basic aspects of mealybugs and their primary endosymbionts. J Evol
of mealybugs and also management tactics Biol 18:315324
known for mealybug species affecting different Franco JC, Suma P, da Silva EB, Blumberg D, Mendel Z
(2004) Management strategies of mealybug pests of
crop plants in different countries. Section I of the
citrus in Mediterranean countries. Phytoparasitica
book presents a generalized description of mor- 32:507522
phology, cytogenetics, taxonomy, molecular Kosztarab M, Kozr F (1988) Scale insects of Central
characterization, biology, damage, ecology, natu- Europe. Dr. W. Junk Publishers, Dordrecht
McKenzie HL (1967) Mealybugs of California. University
ral enemies, ant association, control measures,
of California Press, Berkeley, 525 p
insecticide resistance, pheromones, etc. Section Walton VM, Daane KM, Bentley WJ, Millar JG, Larsen
II deals with management practices of mealybugs TE, Malakar-Kuenen R (2006) Pheromone-based mat-
in different crops. ing disruption of Planococcus cus (Hemiptera:
Pseudococcidae) in California vineyards. J Econ
Entomol 99:12801290
Williams DJ (1985) Australian mealybugs. British
References Museum (Natural History), London, 431 p
Williams DJ (2004) Mealybugs of southern Asia. The
Ben-Dov Y (1994) A systematic catalogue of the mealy- Natural History Museum, Southdene Sdn. Bhd., Kaula
bugs of the world (Insecta: Homoptera: Coccoidea: Lumpur, 896 p
Pseudococcidae and Putoidae) with data on geograph- Williams DJ, Granara de Willink MC (1992) Mealybugs
ical distribution, host plants, biology and economic of Central and South America. CAB International,
importance. Intercept Limited, Andover, 686 p Wallingford, 635 p
Part I
Mealybugs
Morphology
2
M. Mani and C. Shivaraju
surface of labium. The basal segment usually may be long or short. If they are long, they may
possesses three pairs of setae. At the tip of the extend to or slightly beyond the tip of the claws
apical segment, there is a pair of minute setae that and may be either knobbed or setose at the api-
are usually stiff and spine-like, but because of ces. Digitules less than half the claw length are
their small size they are not shown in the accom- usually setose.
panying illustrations. Immediately anterior to
these apical setae on the anterior surface there are Translucent Dots or Pores They occur on the
usually four pairs of subapical setae. On the hind femur and tibia of quite a few mealybugs.
remainder of the medial and apical segments
there are varying numbers of setae, which reach Clypeolabral Shield In some species, an anterior
their greatest numbers in members of the tribe extension to the clypeolabral shield is present.
Allomyrmococcini associated with herdsmen
ants. There are only two pairs of anterior setae on
the posterior surface of the apical segment in sub- 2.3 Abdomen
family Pseudococcinae, whereas there are three
such pairs of setae in the subfamilies Trabutininae, Dorsal Ostioles The most characteristic feature
Rhizoecinae and Sphaerococcinae. One pair of of the family Pseudococcidae is the occurrence of
the subapical setae is grooved on the labium of two pairs of slit-like openings on the body dor-
Phenacoccus manihoti. sum, here designated as dorsal ostioles. The pos-
terior pair lies within the boundaries of the
seventh abdominal segment, and the anterior pair
appears to belong to the foremost part of the pro-
2.2 Thorax thorax. The edges of the ostioles are invaginated
to form anterior and posterior lips, and these are
Spiracles The spiracles in the Pseudococcidae usually beset with setae and trilocular pores.
are represented by two thoracic pairs only. The When a living mealybug is disturbed or irritated,
anterior pair of spiracles is located in the inter- a globule of liquid is often discharged from one
segmental membrane between the prothorax and or more of these ostioles.
the mesothorax. In the same manner, the poste-
rior pair of spiracles indicates the border between
the mesothorax and the metathorax. In a few spe- Cerarii These structures number at most 18
cies of Antonina and certain other grass-infesting basic pairs. A cerarius is often composed of two
forms, the spiracles are noticeably enlarged, or more conical to lanceolate setae and a compact
sclerotised and often have a conspicuous crescent group of trilocular pores. Each cerarius produces
of crowded trilocular-type pores situated around a lateral wax lament when viewed alive. The
the lateral margin of the atrium. Usually, how- number of cerarii may vary even between species
ever, the spiracles are essentially the same size in the same genus or cerarii may be absent
and shape throughout the family. entirely. Sometimes a cerarius may consist of
only a single conical seta or there may be multi-
Legs A principal leg character was considered ple conical setae. In some species, there are addi-
to be the presence or absence of a denticle or tional, intermediate cerarii present and the cerarii
tooth on the plantar surface of the claw. This may appear to form a continuous row, when it is
tooth has, at its very highest development, a quite difcult to determine the number of basic pairs.
insignicant character, yet it correlates very Usually in the second and third instars, the cerarii
closely with other characters, which in their total- are more clearly dened so that the total number
ity dene the genera that may be referred to as the can be veried. In some cerarii, the conical setae
Phenacoccus series. The claws bear two apically may be replaced by agellate setae surrounded
spatulate or setose digitules that arise, one on by trilocular pores, or a cerarius may contain one
each side, from near the claw bases. The digitules conical seta and one agellate seta. It has been a
2 Morphology 9
Generalised and semidiagramatic drawing representing morphological structure of mealybugs (Courtesy: Williams DJ)
10 M. Mani and C. Shivaraju
custom to refer to the rst three cerarii on the Circulus The circulus when present consists of
head as the frontal, preocular and ocular cerarii. a simple, sclerotised ring enclosing an area of
They are numbered from the anterior end down- variable size. It may be situated on the venter in
wards, as are all body characters. Each of the full the intersegmental fold between the fourth and
complement of cerarii is numbered as C118, with fth abdominal segments, or on the fourth
C1118 occurring on abdominal segments I abdominal segment above. It encloses an area
VIII. The segmentation on the thorax is some- which is free from pores and setae.
times not clearly dened but by tracing the lateral
ends of the intersegmental lines when possible, it Pores and Ducts Several different types of pores
seems that there are two cerarii present on each and ducts on the body may be recognised in the
thoracic segment (C510) and four on the head Pseudococcidae, which include bitubular and trit-
(C14). There are many species with 17 pairs of ubular (sometimes called bi- or tritubular cera res),
cerarii, when C2 (the preocular pair) is missing. trilocular, minute circular (sometimes called sim-
When only a single pair of cerarius is present, it ple disc pores), multilocular (sometimes called
is located on the anal lobes only (C18). Some spe- discoid or genacerores) and quinquelocular types.
cies possess additional, dorsal cerarii.
Trilocular Pores or Swirled Pores They are
Anal Ring The anal ring (anal opening) in the usually present in species of the family
Pseudococcidae is situated on what is here inter- Pseudococcidae. Occasionally, they are larger on
preted as the tenth abdominal segment. The anal the dorsum than on the venter and in some spe-
ring usually lies on the dorsal side of the body situ- cies of Rastrococcus, those in and near the cerarii
ated close to the posterior apex of the abdomen. In are different in shape and size to others elsewhere
some cases, it may be displaced anteriorly on the on the body. Some trilocular pores in Antonina
dorsum and lie some little distance from the poste- and Chaetococcus are as deep as they are wide.
rior apex of the abdomen, and in rare instances it
may be displaced posteriorly to the venter. The Discoidal Pores These are usually minute, sim-
anal opening is usually surrounded by a more or ple, circular pores present in varying numbers over
less sclerotised ring that normally bears six or the dorsum and venter. In some species of
more slender setae. In this sclerotised band in most Dysmicoccus, the discoidal pores have a granular
members of Pseudococcidae appears numerous surface. The rim of each pore may be thin or con-
irregular pores. In a few instances, the ring is much spicuously wide and heavily sclerotised. In the
reduced, the sclerotization is slight and the pores genus Stricklandina, there are normal minute
are absent. pores present and others with thick sclerotised
rims and a granular or tessellated surface.
Anal Lobes The anal lobes are situated on the Occasionally, discoidal pores are oval, as in some
more or less protruding posterior areas of the species of Eurycoccus. In Hordeolicoccus, some
ninth abdominal segment. On the ventral surface, species possess remarkably large discoidal pores,
they possess at the apex usually the longest body each about the same size as a multilocular disc
seta, here designated as anal lobe seta. On the pore. An unusual type of discoidal pore is described
dorsal surface of each anal lobe is a cerarius, herein for some species of Exallomochlus, in
probably more prominent than others along the which the centre of the pore is extended.
body margin because of more trilocular pores,
slender auxiliary setae, two to several stout coni- Tubular Ducts There are many variations in the
cal setae and often a sclerotised dorsal surface. tubular ducts. The presence or absence of oral
rim ducts is sometimes difcult to decide
Vulva The presence of vulva is an indication of because, although the rim may be present, it may
full maturity of the adult female. It is important as not be elevated from the surface of the derm, as in
a landmark to indicate the exact position on the Leptococcus species. Sometimes, oral collar
venter of the anterior margin of the ninth and pos- tubular ducts possess indistinct rims, which are
terior margin of the eighth abdominal segments. discussed in this chapter.
2 Morphology 11
Microducts Structures that appear as minute present with two conical setae in the abdominal
dots on the surface of the cuticle are actually segment IX. Normally, one trilocular disc pore is
microducts. They may be common throughout present in the cerarian zone of each segment.
the Pseudococcidae. Head: Six jointed antennae, average measure-
ments of the segments in are I, 22; II, 21; III,
Ostioles Normally, they are present as two pairs 17; IV, 16; V, 16; VI, 55. Eye is about 15 in
and lie submedially on the dorsum but in the diameter at the base and 7 high. Beak is conical,
Allomyrmococcini they are situated on the lateral on an average 62 long and 40 wide at the
margins, when the sclerotised lips are base. Thorax: Average measurements of posterior
prominent. leg (in ) are as follows: trochanter, 30 19;
femur, 68 27; tibia, 55 18; tarsus, 65 15;
Body Setae Most pseudococcids have at least a claw, 16; tarsal digitule, 23; claw digitule, 15.
few small dorsal setae, and some are quite setose. Both anterior and posterior spiracles are about 6
In certain species the setae are very slender, while in diameter at atrium and about 15 long.
in others they may be stout and conical or lanceo- Abdomen: Anal ring is situated in between two
late, often the same size as that in the cerarii. anal lobes, 26 in diameter; anal ring setae are 44
Rarely the stout setae may be truncated apically, long on an average. Apical setae are 135 long
and at times they may be borne upon a sclerotised on an average. Anal lobe bar is weakly sclero-
process. The setae on the venter are usually slen- tised. Dermal structures: Only one trilocular disc
der, and normally are situated in transverse rows pore is present in each lip of both the anterior and
on the abdominal segments, in a group anterior to the posterior pairs of ostioles. Trilocular disc
the clypeus, and on areas designated as sternal in pores, about 3 , are present in transverse rows on
the thoracic segments. Infrequently the setae are
of taxonomic value at the species level.
of posterior leg in are trochanter, 66 34; rax, 23; mesothorax, 36; prothorax, 42; head, 20;
femur, 127 53; tibia, 121 30; tarsus, 87 26; and ventral abdominal segments IX, 4; VIII, 11;
claw, 27; tarsal digitule, 36; claw digitule, 24. VII, 12; VI, 13; V, 10; IV, 10; III, 12; II, 12; meta-
Anterior spiracle is about 34 long and 13 thorax, 15; mesothorax, 28; prothorax, 24; head,
wide at atrium; posterior one is 36 long and 13 13. Tubular ducts of oral rim type are present
wide. Abdomen: Anal lobes are prominent; anal mostly on dorsum and a few on venter. Ducts on
ring on an average is 60 in diameter; anal ring dorsum are 9 long and 6 wide. Ventral ducts
setae are 84 long; apical setae is 209 long on are about 3/4 wide of those in dorsum. Their
an average. Anal lobe bar is moderately sclero- approximate numbers are dorsal abdominal seg-
tised. Dermal structures: Ostioles can be found ments IX, 6; VIII, 11; VII, 4; VI, 10; V, 11; IV, 14;
with a few trilocular disc pores. Anterior pair III, 14; II, 14; metathorax, 21; mesothorax, 27;
with three pores and one seta on each lip; poste- prothorax, 19; head, 7; and ventral abdominal seg-
rior ones with three to four pores and zero to one ments IX, 0; VIII, 2; VII, 2; VI, 2; V, 2; IV, 2; III,
seta on each lip. Dorsal setae are of two sizes, 3; II, 3; metathorax, 2; mesothorax, 6; prothorax,
longer and stout, and shorter and thin. Ventral 6; head, 4. Tubular ducts are of oral collar type,
body setae are longer and agellate. Circulus is 3.54.0 long and 1.5 wide, mostly distributed
about 33 long. Trilocular disc pores, 3.23.6 , in the marginal and submarginal areas of venter,
are present on both the surfaces of the body but rarely found on dorsum. Their approximate num-
more on dorsum. Their approximate numbers are bers in venter are abdominal segments IX, 0; VIII,
dorsal abdominal segments IX, 14; VIII, 15; VII, 2; VII, 4; VI, 5; V, 4; IV, 4; III, 4; II, 7; metathorax,
32; VI, 19; V, 18; IV, 19; III, 30; II, 30; metatho- 10; mesothorax, 8; prothorax, 4; head, 4.
Third-instar female nymph of M. hirsutus (Green). Adult female of M. hirsutus. MLDP multilocular disc
OCD oral collar duct, ORD oral rim duct pore, VUL vulva
(Courtesy: Ghose SK)
14 M. Mani and C. Shivaraju
2.4.4 Adult Female former being predominant in the dorsum and the
latter in the venter. Oral rim ducts of dorsum are
Body is ovoid, slightly broader and rounded at larger than those of venter and more or less
the anterior end, on an average 1.7 mm long and arranged in transverse rows, about 9.5 long and
1.1 mm wide, attaining larger size (3.2 mm l.7 45 in diameter at the opening. Their approxi-
mm) with maturity. Anal lobes are prominent, mate numbers are abdominal segments IX, 12;
particularly in young adults. Six pairs of cerarii VIII, 20; VII, 10; VI, 22; V, 32; VI, 35; III, 34;
in the abdominal segments IVIX, usually with I1, 36; metathorax, 54; mesothorax, 62; protho-
two cerarian setae are present; occasionally a rax, 42; head, 18. A few rim ducts of venter are
third one is present in the cerarii of segments VIII found in the marginal and submarginal regions of
and IX. Segment IV has generally only one cerar- the body. Their numbers are abdominal segments
ian and one stout and longer setae on one side, IX, 2; VIII, 4; VII, 3; VI, 5; V, 5; IV, 7; III, 7; II,
the other cerarius has two normal cerarian setae. 6; rnetathorax, 4; mesothorax, 7; prothorax, 8;
Cerarii are without auxiliary setae except the anal head, 4. Oral collar ducts of venter are variable in
lobe pair. Each cerarius of segment IX has 56 size, 2.42.8 in diameter at opening and on an
trilobular disc pores and three oral rim ducts. average 10.5 long. The ducts of the dorsum are
Head: Antennae appear to be nine jointed generally smaller. These ducts are much more
because of a pseudo-articulation in the terminal numerous (six to seven times) in the adult than in
joint. Average measurements in are I, 54; II, 54; the third nymphal female. Multilocular disc
III, 52; IV, 34; V, 41; VI, 40; VII, 39; VIII, pores, 5 in diameter, are restricted to the sub-
37 + 56. Eye is about 32 wide at the base and 22 marginal and median regions of venter, mainly in
high. Beak is conical, on an average 141 long the abdominal segments VIIX.
and 86 wide at the base.
Thorax: Average measurements of posterior
leg in are trochanter, 97 36; femur, 217 68; 2.4.5 Second-Instar Male Nymph
tibia, 227 32; tarsus, 100 27; claw, 33; tarsal
digitule, 49; claw digitule, 31. Tarsal digitule, of Body is oval with anterior end slightly broader
the anterior legs are unequal, one is about 49 , and rounded; anal lobes are prominent. Average
whereas the other is about 42 . Anterior spiracle body size in the early stage is 625 long and 390
is about 51 long and 26 wide at atrium, and wide. It increases greatly and attains 970 438
posterior one 55 long and 29 wide. at the end of the feeding period. Normally, one
Abdomen: Anal ring on an average is 72 in pair of cerarii present are in the abdominal seg-
diameter; anal ring setae is 154 long; anal lobe ment IX; generally each with two and rarely stout
bar is moderately sclerotised; apical setae are 251 conical setae, one auxiliary seta, one microduct of
on an average. Dermal structures: Anterior pair oral collar type and one trilocular disc pore.
of ostioles with nine to ten trilocular disc pores Segment VIII is occasionally with one or two cer-
and one to three setae on each lip are present; arian setae in each. The cerarian setae of other
posterior one with 912 pores and 14 setae on segments are slender and elongated. Generally,
each lip. Body setae are of two sizes on both dor- two disc pores and one collar duct are present in
sal and ventral surfaces, the ventral ones being each cerarian zone of other segments. Head: Six
generally longer. Circulus is about 77 long. jointed antennae, but the joints cannot be recog-
Trilocular disc pores are more numerous and nised as and when the antennae of third-instar
larger on dorsum, about 4 , whereas those on male nymphs develop inside this instar. Average
venter measure about 3 . These pores are much measurements of the segments in are I, 32; 11,
more numerous (above 20 %) in the adult than in 27; III, 34; IV, 20; V, 22; VI, 62. Eye is about 22
the third-instar females. Tubular ducts are of two in diameter at the base and 10 high. Beak is
types: oral rim ducts and oral collar ducts, the conical, on an average 94 long and 57 wide at
2 Morphology 15
the base. Thorax: Average measurements of pos- imate numbers are dorsal abdominal segments IX,
terior leg in are trochanter, 46 26; femur, 9; VIII, 9; VII, 18; VI, 13; V, 15; IV, 16; III, 22; II,
98 38; tibia, 84 20; tarsus, 74 18; claw, 21; 19; metathorax, 20; mesothorax, 44; prothorax,
tarsal digitule, 34; claw digitule, 20. Anterior spir- 32; head, 26; and ventral abdominal segments IX,
acle is about 29 long and 8 wide at atrium; 4; VIII, 5; VII, 10; VI, 9; V, 8; IV, 7; III, 8; II, 7;
posterior one about 31 long and 9 at atrium. metathorax, 11; mesothorax, 21; prothorax, 16;
Abdomen: Anal ring on an average is 36 in head, 17. The microducts are of oral collar type,
diameter; anal ring setae are 66 long; apical about 7 long, present on both dorsum and ven-
setae are on an average 172 long. Anal lobe bar ter. The ducts in dorsum are wider (3.23.6 )
is moderately sclerotised. Dermal structures: than those in venter about 2.4 ; their numbers are
Three trilocular disc pores and one to three setae dorsal abdominal segments IX, 2; VIII, 5; VII, 2;
on both upper and lower lips of anterior pair of VI, 2; V, 2; IV, 5; III, 3; II, 7; metathorax, 4; meso-
ostioles and two to three pores and zero to one thorax, 6; prothorax, 5; head, 4; and ventral
seta on each lip of the posterior pair are present. abdominal segments IX, 0; VIII, 1; VII, 3; VI, 4;
Circulus is present. Trilocular disc pores are about V, 3; IV, 2; III, 2; II, 2; metathorax, 2; mesothorax,
3 and more numerous on dorsum. Their approx- 2; prothorax, 2; head, 3.
Second-instar male nymph of M. hirsulus. OCD oral Third-instar male nymph of M. hirsutus. ANT antenna, AS
collar duct anterior spiracle, MLDP multilocular disc pore, AO ante-
(Courtesy: Ghose SK) rior ostiole, PO posterior ostiole, AT anal tube, WL
wing-bud
2.4.6 Third-Instar Male Nymph the antennae of fourth-instar male become prom-
inent, as and when these are formed inside the
Body is oval, more rounded at the anterior end, antennae of third instar. Mouthparts are absent.
on an average 1.138 mm long and 0.504 mm Eyes are not discernible.
wide. Sclerotisation is in general very weak. Thorax: Two small wing buds more or less at
Head: Segmentation of antennae is obscure, right angles to the lateral margins of the mesotho-
with the average length being 276 . The joints of rax. Legs are short in comparison with body
16 M. Mani and C. Shivaraju
length, with a few pointed setae. Average mea- present in between the abdominal segments
surements in are trochanter, 52 32; femur, IXX, but its opening on dorsum or venter is not
112 43; segmentation of tibia, tarsus and claw is discernible. Near the posterior end of the abdo-
not well differentiated, their combined length and men, six to seven setae are arranged transversely.
maximum breadth being 175 31. Tarsal and Marginal and submarginal areas of segment IX
claw digitules are absent. Anterior spiracle is dorsally with ve to six more or less transversely
about 29 and posterior one is 29 long and 14 arranged setae are present.
wide. Dermal structures: Both anterior and posterior
Abdomen: Anal ring is absent. A well- pair of ostioles are present. Trilocular disc pores
sclerotised anal tube, 23 long and 26 wide, is are absent.
Fourth-instar male nymph of M. hirsutus. WP wing pad Adult male of M. hirsutus. CS coronal suture, DS
Digitiform seta, QDP quadrilocular disc pore, PS penial
sheath, PW part of wing, PSG penta locular stellate gland,
DE dorsal eye, LE lateral eye, VE ventral eye
2 Morphology 17
Multilocular disc pores: It is 5 in diameter, Dermal structures: Both anterior and posterior
found on both dorsum and venter. Their numbers pairs of ostioles are present, with two multilocu-
are dorsal abdominal segments IX, 1; VIII, 4; lar disc pores and one seta on each lip of posterior
VII, 11; VI, 3; V, 4; IV, 4; III, 9; II, 4; metathorax, pair. Multilocular disc pores, 5 in diameter, are
2; mesothorax, 6; prothorax, 13; head, 8; and present on both dorsum and venter. Their num-
ventral abdominal segments IX, 0; VIII, 2; VII, 3; bers are dorsal abdominal segments IX, 0; VIII,
VI, 3; V, 3; IV, 4; III, 2; II, 2; metathorax, 8; 3; VII, 9; VI, 3; V, 4; IV, 4; III, 4; II, 4; metatho-
mesothorax, 7; prothorax, 6; head 0. Microducts rax, 5; mesothorax, 3; prothorax, 12; head, 0; and
are of oral collar type, about long, present on ventral abdominal segments, IX, 0; VIII, 2; VII,
both dorsum and venter. Ducts of dorsum are 2; VI, 2; V, 2; IV, 3; III, 2; II, 4; metathorax, 5;
about 3.2 wide, whereas those of venter are mesothorax, 5; prothorax, 4; head, 0. Microducts
about 2.4 . Their numbers are dorsal abdominal are of oral collar type, about 7 long, present on
segments IX, 0; VIII, 7; VII, 5; VI, 10; V, 10; IV, both dorsum and venter. The ducts of dorsum are
14; III, 19; II, 19; metathorax, 14; mesothorax, 4; much wider (about 3.2 ) than those of venter
prothorax, 16; head, 4; and ventral abdominal (1.82.4 ) arranged more or less in transverse
segments IX, 0; VI, 3; VII, 4; VI, 7; V, 5; IV, 5; rows. Their approximate numbers are dorsal
III, 8; II, 4; metathorax, 5; mesothorax, 2; protho- abdominal segments IX, 0; VIII, 13; VII, 10; VI,
rax, 6: head, 3. 13; V, 13; IV, 17; III, 17; II, 11; metathorax, 8;
mesothorax, 4; prothorax, 30; head, 4; and ven-
tral abdominal segments IX, 0; VIII, 4; VII, 8; VI,
2.4.7 Fourth-Instar Male Nymph 8; V, 8; IV, 12; III, 01; II, 0; metathorax, 0;
mesothorax, 0; prothorax, 10; head, 0.
Anterior end of the body is round, narrowing Adult males are only of macropterous form,
gradually on the posterior end, on an average on an average 1.055 mm long, including the pro-
1.061 mm long and 0.340 mm wide. Head, thorax jected penial sheath, and 0.310 mm wide. Head:
and abdomen are more differentiated than the Ten jointed antennae, average measurements in
previous instar; sclerotisation is weak. Head: Ten are I, 39; II, 66; III, 79; IV, 69; V, 66; VI, 63; VII,
jointed antennae, average measurements in are 67; VIII, 67; IX, 58; X, 71. The antennae are
I, 34; II, 46; III, 34; IV, 24; V, 27; VI, 29; VII, 32; clothed mainly with digtiform setae, up to about
VIII, 37; IX, 34; X, 74; second segment is the 39 ; a few thicker specialised digtiform setae are
broadest. Mouthparts are absent. Eyes are not present on the last three apical segments, the lon-
discernible. Thorax: Average measurements of gest ones being 39, 49 and 49 on segments VIII,
hind leg in are trochanter; 60 29; femur, IX and X, respectively. Coronal suture is well
128 44; tibia, 142 28; tarsus, 101 25; claw, developed. Dorsomedian sclerite is weakly
16; tarsal and claw digitules are absent. Anterior sclerotised. Three pairs of eyes are present: dor-
spiracle is about 26 long and 13 wide at sal, ventral and lateral. The average diameter of
atrium; posterior one is about 31 long and 16 the dorsal and ventral pairs is 30 and 34 , respec-
at atrium. Wing pads are obliquely attached to tively. Lateral pair is 25 in diameter at the base
the mesothorax. Abdomen: In segment X, six to and 18 high on an average. Mouthparts are
seven setae are transversely arranged on dorsum. absent. Thorax: One pair of wings, on an average
Two marginal setae are on dorsum on each side 0.92 mm long and 0 .42 mm wide; each wing has
of segment IX, the longest one about 63 , and four to ve sensory setae near the basal region;
two corresponding ones on venter about 17 . average measurements of the posterior leg in
Anal tube, apparently without an external open- are trochanter, 62 26; femur, 216 39; tibia,
ing, is present in between segments IX and X, 22 283 23; tarsus, 99 19; claw, 34; tarsal digitule
long and 26 wide. Penial sheath of adult male is very slender, 34. As in antennae, legs are
is visible as and when it is formed inside this clothed with both digitiform and slender-pointed
stage. setae, their maximum length being 31 and 21 ,
18 M. Mani and C. Shivaraju
respectively. The inner distal end of tibia has disc pores are present on each side of the
three spines. Tarsus has three to four spines at the abdominal segment IX. In the centre of each clus-
inner distal end. Anterior and posterior spiracles ter, there are eight to ten disc pores of smaller
are about 21 wide at atrium, their lengths being dimension (about 4 in diameter) and three long
23 and 26 . Abdomen: Penial sheath is about 179 setae, two of which on an average are 260 long.
long and 70 at the widest portion and 6.5 at Around the central zone 3844 disc pores, 5 in
the projected tip, which is rounded. It has two diameter, are present.
distinct median lobes, each more or less triangu-
lar in shape. Dermal structures: Only posterior
pair of ostioles is present. Quadrilocular disc References
pores, 4.85.6 in diameter, are present on both
dorsum and venter; numbers on dorsum are Ghose SK (1971) Morphology of various instars of both
sexes of the mealy-bug, Maconellicoccus hirsutus
abdominal segments IX, 0; VIII, 4; VII, 2; VI, 2;
(Green) (Pseudococcidae: Hemiptera). Indian J Agric
V, 3; IV, 2; III, 2; II, 11; metathorax, 0; mesotho- Sci 41(7):602611
rax, 0; prothorax, 816; head, 4; and ventral McKenzie HL (1967) Morphology and classication.
abdominal segments IX, 0; VIII, 3; VII, 3; VI, 3; Mealybugs of California, University of California
Press, Berkeley, pp 3640
V, 3; IV, 3; III, 0; II, 2; metathorax, 2; mesotho-
Williams DJ (2004) Mealybugs of southern Asia. The
rax, 48; prothorax, 48; head, 0. Two dorsal Natural History Museum, Southdene Sdn. Bhd., Kaula
clusters of stellate or tail-forming pentalocular Lumpur, Malaysia, pp 1821
Cytogenetics
3
Ramakrishna Sompalaym, Kokilamani A.
Lingarajaiah, Raju G. Narayanappa, Jayaprakash,
and Venkatachalaiah Govindaiah
Jayaprakash
Department of Zoology, Bangalore University,
R. Sompalaym (*) K.A. Lingarajaiah
Bengaluru 560 056, India
Department of Zoology, Bangalore University,
Bengaluru 560 056, India Centre for Applied Genetics, Bangalore University,
e-mail: srkbuz@ymail.com Bengaluru 560 056, India
R.G. Narayanappa V. Govindaiah
Department of Biotechnology, KSOU, Centre for Applied Genetics, Bangalore University,
Mukthagangotri, Mysore 570 006, India Bengaluru 560 056, India
been sporadic reports that provide incomplete and during successive divisions. In the absence of
contradictory information pertaining to Indian kinetochore-based cell divisions that prevail in
fauna for their chromosome systems (Tulsyan coccoid chromosome systems, and also, in order
1963; Dikshith 1964, 1966; Chauhan 1970, 1977). to accommodate the occurrence of karyotypic
changes, Brown (1961) assays chromosome frac-
ture and fusions in the place of the prevalent
3.2 Mealybug Chromosomes nature of chromosomal rearrangements. It was
also envisaged that simple breakage can deter-
All coccoids possess holocentric chromosomes, mine increase or decrease in chromosome num-
that is, diffuse centromeres (Hughes-Schrader bers unless a breakagefusionbridge cycle
and Ris 1941). Inverse meiosis is a second ances- intervenes to eliminate the breakage points.
tral condition manifested in coccoids that is also Species relationships can be explained by cit-
shared with the other closely allied aphids (Ris ing chromosome variability occurring with
1942; Hughes-Schrader 1944, 1948). respect to either chromosome numbers or mor-
Although the cell cycle sequence is different phology. Brown (1961) insists upon spontaneous
from that of typical meiosis, results are the same; occurrence of chromosome breakage resulting in
each of the four chromatids of meiotic bivalents abundant availability of ruptured chromosomes
reaches one of the four nuclei produced by meio- for increase in the diploid numbers either by
sis. Coccoids are also manifested by those systems chance or incurred by selection. There are an
in which at least some of the females are produced abundant number of cases dealing with karyo-
parthenogenetically, in addition to the usual bisex- typic changes incurring based on chromosome
ual mode of reproduction. They are considered to fragmentation in mealybug genomes (Nur et al.
have unique chromosome systems and they offer 1987; Cook 2000).
enormous potential in our understanding of prob- Changes in chromosome numbers with respect
lems such as chromosome imprinting and differ- to pseudococcid species have been reported to be
ential regulation of homologous chromosome sets in the range of 8 to 64 and that ranges within coc-
(Chandra 1971; Chandra and Brown 1973). coids are rather small compared to other insect
Chandra (1971) suggested for the rst time that groups (Hughes-Schrader 1948; Nur et al. 1987).
there are some similarities and also contrasts Until now, 115 cytogenetically studied species of
between mammalian X-chromosome inactivation mealybugs belonging to 44 genera have been
and the inactivation of paternal chromosomes in made known (Gavrilov 2007; Gavrilov and
mealybugs. These include genomic imprinting, Trapeznikova 2007, 2010). Eventhough, the dip-
facultative heterochromatization, and differential loid number of chromosomes ranges from 8 to 64,
regulation of homologous chromosomes. the modal number seem to fall on 10 (Plate 3.6
Subsequently, Brown and Chandra (1977) have Fig. 4, 6, 8). Few mealybugs showed intrageneric
drawn attention to emphasize that coccoids are at variation in their chromosome numbers; for exam-
the pinnacle of an evolutionary pyramid of cytoge- ple, in genera such as Antonina (2n = 12, 16,
netic variants and complexity. In order to under- 24 + Bs), Nesopedronia (2n = 18, 14, 10) and
stand these variations in chromosome mechanics, Trionymus (2n = 16, 10, 8), such instances can be
it becomes essential briey to review pseudococ- cited as useful in taxonomic and phylogenetic con-
cid chromosomes. siderations of the genus. Accessory chromosomal
elements (B-chromosomes) were found in several
species of mealybugs (Nur et al. 1987; Gavrilov
3.3 Chromosome Numbers 2004). But, the detailed investigation of
and Chromosome Forms B-chromosomes has been done only in
Pseudococcus viburni (Signoret; Nur 1962a,
Coccoid chromosomes lack specied centro- 1966a, b). The majority of pseudococcids possess
meric regions. It appears obvious to point out that 2n = 10 (Nur et al. 1987; Moharana 1990; Nur
chromosome fragments perpetuate themselves 1990; Gavrilov and Trapeznikova 2007, 2010).
3 Cytogenetics 21
Excepting Planococcus citri and a few other spe- species. Based upon female pachytene chromo-
cies, the number of species studied based on meric sequences, Raju (1994) made an initial
employing recently evolved cytogenetic tech- attempt to describe karyotype and comparison of
niques is very low. One of the reasons cited was three species of the Indian genus Planococcus
the difculties incurred in procuring enough cells (viz. P. citri, P. lilacinus and P. pacicus) essen-
for the preparation of chromosomes and of under- tially based on differential banding patterns, but
standing of chromosome basics for detailed cyto- was unable to identify individualistic karyotypes
logical analyses. For cytological investigations of because of lack of discriminating cytogenetic fea-
Indian mealybug taxa, Parida and Moharana tures (Plates 15). Gavrilov (2004a, 2007) and
(1982) and Moharana (1990) attempted to enu- Gavrilov and Trapeznikova (2007, 2010) have
merate chromosome numbers based on conven- made elaborate studies resulting in the elucidation
tional cytological techniques and they were also of the karyotype for more than 25 species of
able to present preliminary assessments of karyo- Russian mealybugs based on squashing techniques
morphological features for more than 20 different for chromosomal preparations. Nur et al. (1987)
were able to describe the karyotype of about 80 Mckenzie 1967 (California), Drozdovskiy 1966
different species of mealybugs that were collected (Russia), Brown 1961 and Hughes-Schrader 1935
from various parts of Africa, America, and a few (USA), and Schrader 1923a (USA) have contributed
from South Asia. Tremblay et al. 1977 (Italy), enormously to the eld of mealybug cytogenetics
3 Cytogenetics 23
in the form of karyological studies. In an attempt complements did not provide any discriminative
to analyse mealybug chromosome morphology, cytological signatures other than suggesting that
chromosome preparations were studied through they belong to and qualify themselves as belong-
the uorescent microscopy using appropriate dyes ing to the Lecanoid type of chromosome system
(e.g., Quinacrine Mustard (QM)/QM dihydrochlo- (Jaipuriar et al. 1985; Venkatachalaiah and
ride), and it was found that these chromosomal Chowdaiah 1987; Venkatachalaiah 1989).
24 R. Sompalaym et al.
3.4 Telomeres and C- Bands C-staining proles along the length of each chro-
mosomal fragment in the complement.
It is of interest to note that with particular impor- Employing classical C-staining protocol upon
tance to the diffuse nature of centromeric systems Planococcus citri metaphase chromosomal prep-
manifested by coccoid chromosome morphology arations, it was expected to highlight constitu-
it was expected to display discriminative tively heterochromatic sites in the complement.
3 Cytogenetics 25
But the cursory observations led in demonstrat- this chromosomal component may offer convey-
ing that the C-specic bands were found speci- ing information about its cytogenetic context.
cally identifying the telomeric region specicity The situation acquires a genetic signature due to
in the metaphasic chromosomal complement and its co-orientation pairings during late meiotic
this situation was ascribed as T-bands (male or female) chromosome synaptic processes
(Venkatachalaiah 1989; Raju 1994). However, (Plate 3.7 Fig. 6, 7, 8, 9).
the results obtained by Venkatachalaiah (1989) Ferraro et al. (1998), in their attempt to local-
and Raju (1994) pertaining to C-banded stainings ize C-banded regions at P. citri chromosomal
at telomeric ends of each chromosome in the preparations, found evidence regarding
complement were irrespective of a particular C-positive bands localizing at the telomeric
chromosome type (whether of mitotic, meiotic, regions of all chromosomes in the complement.
or polyploid nuclei) or sexes (males or females), When they further insisted upon prior exposure
and thus, they contend that these cytological to CMA3 (chromomycin A3) -methyl green and
markers could be representing a particular type subsequent exposure to C-staining protocols, the
of constitutive heterochromatic component. The implicit C-bands were found correspond to telo-
intense stainability at the telomeric regions in the meric region-specic areas. This has led them to
chromosomal content allows one to assay that infer that these results could, however, representing
3 Cytogenetics 27
Plate 3.7 Representatives of other pseudococcids showing NOR- entities and C-band regions
GC-rich specic spots on chromosomes. From the point of view of cytology, telomeres
However, when they insisted further upon are marked by specialized DNA and protein
H33258 uorochrome to live cells prior to components that usually decorate the chromo-
C-staining, they recovered images almost imitat- some ends or other specic loci. In several
ing C-banded telomeric-specic regions. Thus eukaryotes, their occurrence and prevalence has
they were able to interpret the failure to nd a been tested, wherein they have been found com-
dull appearance, instead of bright bands at the posed of simple tandem pentameric (TTAGG)
specied locales, and those of brighter and repeats localizable at specic chromosome loci
intense bandings could represent condensed con- accompanied by complex subtelomeric struc-
stitutive heterochromatic regions, leading them tures in close apposition (DAiuto et al. 2003; De
to insist that there could be more DNA congre- Lange 2005). A large number of molecular cyto-
gated per unit length per chromosome than in the logical studies have led to the implication that
euchromatic zones. Moreover, some of the telo- telomeres of eukaryotes are usually composed of
meric regions being positive to DAPI stainings, it conserved short tandemly repeated GC-rich
was inferred that the presence of AT-rich sequences. This kind of sequence conservation is
sequences were embedded within the predomi- reected as a common mechanism for telomere
nantly GC-rich regions of individual region biosynthesis. This mechanism specically
chromosomes. dictates and involves the activity pattern of a
28 R. Sompalaym et al.
kinds of cytological sites, and thus probably rep- (Nucleolar Organizer Region) specicities on
resented by either telomeric and/or centromeric metaphase chromosomes. These results point to
sequences. Schweizer and Loidl (1987) have pro- have driven them to ascribe that the FISH tech-
posed a model that explains how nique might help in identify with P. citri chromo-
C-heterochromatin enhances and leads to adher- somes at specic zones on all chromosomes
ence of such chromosomal zones conning and/ except at one pair in the complement. But silver
or inducing towards effecting interchanges of nitrate staining specicity had enabled in speci-
heterochromatic material between nonhomolo- fying at a single pair in the complement but char-
gous and homologous chromosomes in the com- acteristically demonstrating the site at which
plement and thus leading towards annealing into bearing very prominent macer-shaped, silver
a common platform resulting in such situation nitrate stainable entities, irrespective of their ori-
that they belong to as though in a monocentric gin whether of euchromatic or heterochromatic
type of chromosome system; that also insists chromosomal pair (Plate 3.7 Fig. 1, 2, 3, 4, 5).
upon those of chromosomal regions with holoki-
netic activity that do not t into this model. In
view of the limited information gathered from 3.5 B- Chromosomes
other homopteran examples, an effort was made
to dene that the nature and kind of telomeric During the courses of systematic cytological
components that were found enhanced to estab- study in the case of Pseudococcus afnis chro-
lish as a C-banded heterochromatin. Moreover, mosomal complement that possesses supernu-
Panzera et al. (1992) and Prez et al. (1997) merary B chromosomes which were transmitted
based on their limited experience offer the opin- without the reduction during spermatogenetic
ion, especially of Triatoma meiotic systems, that courses that were found exhibiting a strong mei-
the tendency of the heterochromatin component otic drive, in such processes (Nur 1962a, b,
inferring to change in accordance with from one 1969). Prior to spermatogenesis, the B chromo-
chromosome to another or from proximal to dis- some was heterochromatic, but during prophase I
tal sites of the same chromosomes within a com- of spermatogenesis it became evident that even
plement. However, this characteristic cytological less condensed than the euchromatic set (i.e.,
feature was found on preferential localization of negatively heteropycnotic) and this change in
telomeric heterochromatic content of some condensation property apparently makes this sit-
instance cases alone probably thereby reecting uation possible for the Bs to segregate with the
upon C-banded components. These proposals are euchromatic set and be transmitted over to 90 %
in congruence with those of the Schweizer and of the offspring. Nur and Brett (1985, 1987,
Loidl (1987) hypothesis, but this type of chromo- 1988) have presented subjective data supporting
somal behavior is not in any way agreeable to that acquisition of the condensation property of
certain terms with other instance cases analyzed As and Bs during spermatogenesis seemed to
from other homopteran examples for the said infer that this situation is due to the presence of
purposes including the coccoid chromosome genotype that affects the rate of transmission of
systems. the Bs in males. However, it is somewhat clear
Ferraro et al. (1998) had undertaken an elobo- that this situation became evident because of the
rate proceedings in view of eliciting and appro- inuence of this genotype which has affected the
priating the preponderance of the ribosomal condensation property of B, but not the property
cistrons and upon highlighting of their cytologi- of heterochromatization. However, Klein and
cal localization based on the mealybug chromo- Eckhart (1976) theorized that difference between
somal preparations. This analysis had led to the Bs and regular chromosomes of Pseudococcus
results so obtained by means of the FISH tech- afnis could be due to changes occurring at the
nique and of subsequently staining the same with DNA sequences level. Another probable reason
silver nitrate solution for localizing NOR sighted was the differences observed between the
30 R. Sompalaym et al.
A and B sets that could be due to the occurrence licate or having replicated once, the euchromatic
of DNA of the two types of heterochromatin that sets replicated several times (Lorick 1970; Nur
being methylated. Thus, the percentage of 1966c, 1970, 1972).
5-methylcytosine in the DNA of P. afnis was The sex-specic association of the microor-
found to be higher in males than in females, and ganisms has led to the suggestion that they may
higher in females without Bs than in females with have a role in sex determination (Buchner 1965).
Bs (Scarbrough et al. 1984). However, the precise nature and role of endo-
symbionts in normal development has not been
clearly assessed. Biochemical and morphological
3.6 Polyploidy analyses of isolated endosymbionts have estab-
and Endosymbionts lished their prokaryotic characteristics (Houk and
Grifths 1980; Ishikawa 1989). The 16 s rRNA
In most species of mealybugs the polar bodies re- gene sequences of several homopteran insect
enter the egg and contribute to or give rise to endosymbionts including those of certain species
large polyploid cells (mycetocytes) that house of mealybugs and aphids, have been considered
intracellular bacterial symbionts (Brown 1965). for their role in the prevalence of phylogenetic
In some mealybugs, cells formed by polar bodies relationships among those species probed for
1 and 2 are known to be totipotent. In male those purposes (Munson et al. 1991, 1992;
mealybugs and in other coccoid families, one Kantheti 1994). However, the nature and extent
portion of the genome becomes heterochromatic of type of expression of the concerned gene
and the other becomes euchromatic (genetically inquisition during the course of insect develop-
active) in several tissues or organs (Tremblay ment are not clearly explained. Buchner (1965)
and Caltagirone 1973). These include the midgut, reported that extracellular symbionts are present
the Malpighian tubules, the salivary glands, in the females of Stictococcus but absent in the
oenocytes, and serosa (Nur 1967, 1972). One males. Most coccoids contain intracellular bacte-
characteristic of most of these tissues is that their ria or yeastlike symbionts present in the cyto-
nuclei later become polyploid as a result of endo- plasm of special cells, the mycetocytes (Tremblay
reduplication or endomitosis (Plate 3.6 Fig. 3, 5, 1977, 1989). The origin of the mycetocytes is of
9). During oogenesis, polar bodies do not degen- interest because it may vary between, as well as
erate; instead they re-enter the egg cell, and fuse within, families. Therefore, it appears probable
with each other and also with some of the cleav- that the origin of mycetocytes may have an
age nuclei and form polyploid cells called myce- important bearing on the pseudococcid genetic
tocytes. These mycetocytes are invaded by system (Hughes-Schrader 1948).
certain maternally transmitted microorganisms Interestingly, Kantheti et al. (1996) reported
generally referred as symbionts. Mycetocytes an isolation of the 16S rRNA gene sequenced
harboring such symbionts form an organ called segment, designated as P7 from an embryonic
mycetomes whose function is not known (Brown cDNA library of Planococcus lilacinus, which
1965; Nur 1977). Such symbionts are transovari- was found to be an encouraging attempt and by
ally transmitted to the next generation and thus hybridizing to the genomic DNA of females to
show maternal inheritance (Buchner 1965). the assay, but not to that of males. Interestingly
Euchromatization, however, is apparently not an P7 showed no hybridization to nuclei of either
essential step in the development of these tissues, sex, raising the possibility that it was extrachro-
because these types of tissues involved may vary mosomal in origin. Using electron microscopic
between congeneric species. Moreover, the fre- images, especially of P7 clones but not of P3,
quency of cells in which euchromatization occurs annealing was found to the adult female abdomi-
sometimes varies between individuals. However, nal organ called mycetomes. Electron micros-
in those nuclei in which the paternal genome copy has disclosed the presence of symbionts
remained heterochromatic, it usually did not rep- within the mycetocytes. Sequence analysis
3 Cytogenetics 31
showed that P7 is a 16 s rRNA gene conrming These bizarre genetic systems are of immense
its prokaryotic origin. P7 expression is detectable help in our attempt to understand further upon the
in young embryos of both sexes but the absence occurrence of a variety of sex-determining mech-
of P7 in third instar and adult males suggests that anisms prevailing in scale insects in the light of
the designated gene containing isolated gene their inherent property of inverse meiosis effec-
sequences assay and hence, consideration of pro- tively driving them through the efforts of holoki-
visional endosymbionts are the subject and object netic chromosome mechanics. Most species of
of sex-specic elimination/acquisition type of coccoids are bisexuals with extreme sexual
operating processes. dimorphism but due to precariousness of male
populations at times, some of them have become
parthenogenetic. These complex genetic systems
3.7 Mechanism of Sex appear invigorating due to the involvement of
Determination both the bisexual as well as the parthenogenetic
mode of reproduction. Another noteworthy fea-
In many species, sex determination is associated ture is inicted on them due to the deliverance of
with the inheritance of a heteromorphic chromo- quadrinucleate spermatid formation in many
some pair in one sex. However, not all species mealybug (bisexual) chromosome systems. It is
have evolved from a common ancestor that pos- thus possible to surmise that the various types of
sessed such an heteromorphic sex chromosome meiosis that were confronted within the scale
set. Rather, XXXY sex determination appears insect examples could have arisen in a derivative
to have arisen independently many times in evo- form or in a succeeding form from that of primi-
lution from the XXXO type form. The XXXY tive homopteran (aphidcoccid line) examples
sex chromosomes of ies and mammals also including aphid chromosome systems (XXXO
arose independently, but, the underlying mecha- system). It is thus possible to note that during the
nisms of sex determination are quite different and derivation processes it became inevitable in view
difcult to predict except in molecular terms. of the penchant situation prevailing with those
Coccoids are a unique and very peculiar group participants driving in through to the equatorial
of insects in view of their possessing a highly orientation of meiotic bivalents at rst meiosis
variable mode of sex-determining mechanisms. and of the preponderance of prereduction at
This situation becomes evident through the chiasma.
course of studying complex meiotic processes
incurred in a few select examples analyzed thus
far. Thus, this situation has led to the creation of 3.8 XXXO System
some academic interest by some earlier cytolo-
gists to pursue further upon attempting under- Sex determination in primitive coccoids could
stand the intricacies of meiosis and mitosis. have taken its initiation based on the XX ()XO
Interestingly, White (1973, 1978) took special () type of sex-determination mechanism.
interest in accommodating this opportunistic sit- Consequent upon this exigency, oogenesis is of
uation prevailing in mealybugs (scale insects) conventional type progressing through inverse
summarily termed as aberrant genetic systems, meiotic pathways, whereas spermatogenesis is
and Nur (1980) proclaimed unusual chromo- highly modied in most coccoids, mealybugs in
some systems but recent views indict them particular. In view of this unique situation, vari-
either as the more diverse or asymmetric able modes of expression pathways become
genetic system. Serendipity, as applied to these imminent as represented among analyzed primi-
scale insects, which are characterized by posses- tive margarodid examples. Currently, cytological
sion of a peculiar genetic system, was not found records have become known from margarodid
in any other animal system of comparable nature. assemblage of species that include taxa belong-
32 R. Sompalaym et al.
ing to Margarodidae, Ortheziidae, and Putoidae. cies of Puto demonstrated conventional meiotic
The cytological descriptions of these primitive chromosomal features. These species follow a
groups of species characteristically reveal that typical heterogametic mode of sex-determination
the sex of progeny is predetermined prior to and mechanisms, in which the males usually possess
at spermatogenesis; spermatozoa with the one chromosome less than that of females; char-
X-chromosome produce females and spermato- acteristically the example includes Puto species,
zoa lacking it produce males (Nur 1980). In this 2n = 1413; Puto albicans, 2n = 2019; and
extent, Brown (1977) placed primary emphasis Callipappus rubigonosus, 2n = 1413 (Brown
upon the coccoid chromosome system imparting and Cleveland 1968; Hughes-Schrader 1944).
the implicit nature of acquiring adaptive special- There are other taxa in which spermatogenesis is
ization and the same was found reected in the highly modied as was shown in meiosis of
progression of meiotic processes which in turn Protortonia and Matsucoccus gallicola dening
enabled categorizing into three types: (1) multiple sex chromosome systems. Surprisingly,
Margarodid assemblage, (2) Lecanoid types, and the only other report in which no morphologi-
(3) Diaspidid systems (Plate 3.8). In some cally identiable sex chromosomes were shown
Margarodid examples studied, spermatogenesis is represented by an example showing cytologi-
resembles that of conventional oogenesis, as is cal features for the whole Ortheziidae family,
especially evident in the case of Puto. Some spe- comprising 2n = 16 in both sexes (Brown 1958).
3.9 Lecanoid System (i.e., imprinting) of and the activity status (of het-
erochromatization), of which the earlier process
In the Lecanoid chromosome type of coccoids seems likely to be an effective one at the ancestral
that exhibit a peculiar situation among coccoid stock and this attribute might reect in bringing
chromosome systems is one in which one hap- about differential condensation activity of the con-
loid set of chromosomes acquires heteropycno- cerned chromosomes. At this juncture, Brown (1977)
sis during the developmental course and also in felt that this situation appeared premature to the-
the germline cells. Those cells destined to become orize about or make any generalization unless
males acquire this status probably from mid- substantial molecular data were made available.
blastula onwards and persist through to adult life.
In the females, both sets of chromosomes in the
cell remain in the euchromatic state throughout 3.9.1 Parthenogenesis (Unisexual
the developmental course (Schrader 1921, 1923b; Reproduction)
Schrader and Hughes-Schrader 1926). The basis
for such an extraordinary situation in the mealy- While further pursuing the nature of the evolu-
bug chromosome is in the procession of acquisi- tionary trend involved during the course of sexu-
tion of the mechanism of heteropycnosis in male ality of scale insects, Hughes-Schrader (1948)
embryos and this involvement facilitates func- asserted that the prevalence of the parthenoge-
tional inequalities with respect to males that may netic mode of reproduction could be due to con-
point towards proclaiming them as physiological cordance with a higher incidence of disparity in
haploids, even though they have a duplex set in their reproductive potential. Thus, Hughes-
their nuclei (Plate 3.6). Brown and his group Schrader (1948) was able to discriminate bisexu-
have ventured into delineating the processes and als from those parthenogenetic ones and further
involvement of genetic mechanics of genomic suggested the appraisal of three fundamental
inactivation and of cytogenetics of heterochro- types of parthenogenetic products in them. Since
matinization processes in the genome (examples then, there have been considerable amounts of
include Planococcus, Phenacoccus, coccoid cytogenetic information procured by
Maconellicoccus, etc. of the family Brown and his associates that also afrmed that
Pseudococcidae and Laccifer of Kerridae). It is this situation based on cytogenetic surveillance
of interest to learn more about and probe further which acquired an innovative stimulus and pros-
the processes of heteropycnosis of the paternal pects including overall frequency of both bisex-
composition of the Lecanoid chromosomes and ual and parthenogenetic life cycle analysis from
as such it becomes imperative to note that this set among the select taxa of Coccoidea (Schrader
passed through the male phase but expressivity 1923b, 1931; Brown and Bennett 1957; Brown
was conned only to genetic male zygotes. 1963, 1964, 1965, 1966; Nur 1963, 1967, 1969;
Brown and Nur (1964) demonstrated earlier Hartl and Brown 1970). Subsequently, White
through their hybridization experiments that in (1973), who placed greater emphasis upon the
hybrid male embryos the mechanisms of hetero- parthenogenetic mode of reproduction and fur-
chromatinization of the paternal set can occur in thermore, on the implication and validity of het-
the cytoplasm of the foreign cell and thus this erosis during the courses of haplo-diploidy to
mechanism is not neccessarily a species-specic diplo-diploidy, was a matter of great antiquity.
characteristic, because heterochromatinization This and other cumulative studies (White 1978)
progression processes occur after several divi- have moved towards arriving at a conceptualiza-
sions of cleavage, and the paternal set must tion relating to efciency of homozygosity that
somehow seemed to have been marked (or would more likely to have an effective impact
learned) which could have been done or did prior upon haplo-diploids rather than diplo-diploidy.
to the entry of sperm into the egg (Chandra and Brown (1977), Hughes-Schrader (1948), and Nur
Brown 1975). Of the two processes, the marking (1971) have drawn inclinations towards suggesting
34 R. Sompalaym et al.
that it was at the cost of fragility and precarious- mated to have actively participated about eight
ness of males and of their implicit nature of effec- times during the course of evolutionary history
tiveness upon population measures, thereby (Brown 1965; White 1973). It was also felt that
imposing greater inconvenience on the part of the frequency of males (through Arrhenotoky) in
life-cycle strategies. Consequently, this kind of such populations was determined by the fre-
adaptiveness could have been driven towards an quency of haplo-diploids that arose in such con-
alternative mode of reproduction. Thus, the par- siderations. It is thus characteristic of any group
thenogenetic modes could be initiated by means with haplo-diploidy becoming much more
of adapting and involving either of Arrhenotoky inclined towards responding to the oppressive
or Deuterotoky or Thelytoky. But it was at the impact of environmental factors that accrued in
greater behest of Nurs (1969, 1970) concerted which sex ratio potential was deemed to have
efforts that had enabled him in eliciting and cat- been highly variable and it was also found to dif-
egorizing parthenogenomes each exhibiting dis- fer from species to species and even to the extent
tinct types of expression pattern. Subsequently, of genetic strains or of population level extremes.
Nur (1980) was also instrumental in document-
ing a revised format for parthenogenetic modes
of expression based on the following strategies: 3.11 Sex-Ratio Potential
whether the unfertilized eggs develop into males
(Arrhenotoky); or females (Thelytoky) or both In a majority of animal systems studied for the
(Deuterotoky); whether the males are haploid or prevalence of genetic-based sex-determining
diploid; and whether rst meiotic products mechanisms the extent was revealed of separate
between bivalents and oogonia remain the same sexes that direct each whether to become male or
(Gonoid thelytoky) or different (agonoid female, whereas in some taxa, hermaphroditism
thelytoky). may serve the primary mode of reproduction.
Whether a homomorphic (XXXX) or hetero-
morphic (XXXO) mode of sex determination
3.10 Arrhenotoky prevails in them, the sex ratio proportion seems
to be maintained in a harmonious manner (in a
Arrhenotoky is also called as haplo-diploid or 1:1 ratio). In such cases, where conventional dip-
haploid parthenogenesis in which males arise loidy exists, fathers and mothers often obtain
from unfertilized eggs. Males of haplo-diploids equal tness potential through to their sons and
may be referred to as impaternate because they daughters and hence, no sexual conict. But in
have no fathers. From the classical genetic point the case of alternative genetic systems, it becomes
of view, haplo-diploid species may have been essential to involve Trivers and Hares hypothe-
involved in recombinational processes (and sis (1976) that advocates the probabilities pre-
hence, Mendelian in character) inasmuch as they vailing for reproductive success of sons and
behave much to the same extent as those of sex- daughters that can differ markedly from parents.
limited characteristics, but possess no To that extent, they made a proposal in which
Y-chromosome. This, in a way, projects as a sort males and females were drawn into evolutionary
of male heterogamety in genetic characteristics; forces over the aspects of sex-allocation theory
on the other hand, Thelytoky offers a non- that depends on the inclusion and involvement of
Mendelian material thereby propelling it as a a particular genetic system (e.g., haplo-diploidy
reproductive devise. Haplo-diploidy or paternal genome elimination). In such
(Arrhenotoky), on the other hand, is a method of instances, a different set of reproductive trends
sex determination as well as a reproductive sys- seemed to follow in compliance with any biased
tem that involves replacement of an original sex- genetic transmission event that ensues which in
determining mechanism by an entirely new one turn can offer scope for the eventuality of sexual
under extraordinary circumstances. Hence, it conict. With particular reference to mealybug
could have occurred rarely in nature and was esti- examples, it becomes imperative to address the
3 Cytogenetics 35
DusingFischer formula (1976) that insists upon lies in the female germline tissues, suggesting
tness consequences of male and female off- that paternally inherited genes may still have the
spring that can vary with respect to the direct ability to inuence the fate of paternal chromo-
inuence or under duress of genetic and/or envi- somes in the germline but not in soma.
ronmental factors, even though selection prefers Scale insects consequently exhibit consider-
operating in such a way as to bring each into an able variations in their expression patterns based
equilibrium state. These generalizations can evi- on genetic and sex-determining mechanisms, in
dently be tested upon certain cases wherein genes spite of exhibiting similarity in their life-style
in fathers are only transmitted through daughters, strategies (Gullan and Kosztarab 1997; Nur
with sons being of no reproductive potential to 1980; Ross et al. 2011). Due to compulsions of
males. On the other hand, females gain tness their adaptive specializations imposed upon the
benets through both sons and daughters thereby morphology of male and female coccoids, how-
on demand for a possible expression of conicts ever, they differ enormously in terms of certain
over sex ratio. Orienting primarily towards other anatomical features. As is well-known
genetic consequences, the conventional diplo- among scale insects, males are winged hence
diploidy (XXXX system), becomes appar- motile, but with fragile stature although they
ent, in which case they still attempt to maintain a have a short life span based on acquisition of no
rigid sex ratio in the form of 1:1 because mothers feeding habituation, which may eventually suc-
and fathers do not vary through to the contribu- cumb to shortage of male populations. In con-
tions of daughters and sons and hence no sexual trast, females are robust, ornamental, and
conict. However, in unusual situations pertain- sedentary in habituation but have a longer life
ing to scale insects either haplo-diploidy or pater- span and are engrossed with gluttonous feeding
nal genome elimination (PGE) are offered as habits that might propel them to do better in con-
interesting but in extremity for an eventuality (as trolling sex-ratio propensities (Bull 1983).
the case may be). Earlier, Hughes-Schrader (1948) predicted that
On the other side, scale insects exhibit a dif- even though scale insects are besieged with vari-
ferent array of genetic systems, including haplo- able life-cycle strategies leading towards variable
diploidy as well as PGE offering as extremities. modes of genetic schemes, impulsively adaptive
The case of the mealybug (example Planococcus specialization could have driven them to acquir-
citri), provides an ideal system to pursue and ing Thelytoky and hermaphroditism. In fact
probe the kind of involvement and promotion of imposition of such kind of dwellings could have
sexual conict that it exhibits. It has PGE wherein been forced to serve as a clever device to dis-
the male component is in possession of haploid pense with the shortage of males. However,
nuclei, is (either it heterochromatized or) elimi- Brown (1977) contends that in spite of the com-
nated from meiotic cell lineage, and it is present plexities of several chromosome systems within
in somatic cells but untranscribed (Nur 1980). In the scale insects genetic systems, he contem-
terms of genetic mechanisms, the role of genomic plates acquisition of adaptive specialization, in
imprinting may be crucial. Scale insects are turn expanding towards acquiring exponential
known to represent the case in point of genomic taxonomic diversications. However, Nur (1980)
imprinting and imprinting of a paternal chromo- asserts that the fragility of the males may serve as
somal set alone is affected and also acts as a a primary instrument in an easing-out progres-
marker system for sex-determination mecha- sion during the courses of acquisition of a par-
nisms. In the case of mealybug paternal chromo- ticular mode of life-style activities or by adapting
somes especially of Lecanoids it is essential to to a particular chromosome system in succession.
point out that they remain in a latent state during The results obtained based on P. citri, have driven
the course of cell lineages. However, they get James (1937, 1938) and Nelson-Rees (1960) to
transmitted at the cost of a selective advantage. address that in Lecanoids, it appears possible to
Intriguingly, in P. citri, the site of genomic ascribe that females might play an impressive
imprinting of the paternal set of chromosomes role in selection and maintenance of sex-ratio
36 R. Sompalaym et al.
variability by their so adapting towards changes females prior to mating (James 1938; Ross et al.
and at times realigning to overcome any shortage 2010a). An increased understanding of sex allo-
of males. Changes in the sex ratio result in rein- cation theory in mealybugs might therefore yield
forcement of certain changes in developmental insightful opportunities to probe further into the
phases in the females which probably would potential ramications that drive in eliciting evo-
make arrangements towards shifting in the pro- lutionary advantages operating in proceeding
portion of procuring the requisite number of eggs towards extraordinary modes of sex-
and further upon imposing and resetting the determination mechanisms. The results obtained
paternal component onto the gambit of hetero- by Ross et al. (2011) upon P. citri experimenta-
chromatinization drive and in addition in an tions seem to point out appropriate levels that
effort to furthering towards a certain proportion were maintainable based on the role of high tem-
of eggs to be obtained profusely or curtailment. perature, older age at matings, and the starvation
All these adjustments mean amending changes in level, all of which seem to impress during the
contemplation within the scope of remodeling course of the consideration of sex-allocation the-
themselves towards acquiring and procuring a ories. These results may have inuenced changes
sufcient number of males. of expression patterns of female-biased sex
Consideration of imparting environmental ratios. But, they also propose that the effect of
forces upon such forces is based on the popula- temperature seemed rather weak and upon the
tion structure of offspring in responding to tness inuences of food restriction could have strongly
potential which would otherwise be driven implicated in reduced longevity and a transaction
towards differential expression patterns based on of the unusual schedule of male and offspring
the part of engrossing of established conict production across a female reproductive
drawn between parents and offspring. It is known lifetime.
that natural selection operates on those ratios in
which propelling forces rest on whether the par-
ents or offspring are in driving mode (Shuker 3.12 Recent Innovation Made
et al. 2009). It is also suspected that the offspring in Mealybug Genomes
can manipulate the sex-ratio potential thereby
affecting the sex allocation pattern. 3.12.1 Some Molecular Features
Trivers and Willard (1973) have conceived of
a pertinent opinion that environmental factors Heitzs (1928, 1929) unleashing of the opera-
could present oppressive effects on parents with tional denition of the term heterochromatin in
the possibilities of parental interference in an terms of its role in cell-cycle progression has trig-
effort to adjust sex allocation efcacy in the sex- gered momentum in cell biology to roll on
ratio potential. Environmental conditions experi- towards its own toll. This situation came as a
enced by parents can have direct interference natural ingredient for Brown (1966) to elicit suc-
during the course of sex allocation decisions pos- cinctly the ubiquitous nature of heterochromati-
sibly in one of two ways: either directly inuenc- nization serving as a pillar to the cytogenetic
ing parental conditions, or indirectly maintaining conundrum. While nurturing functional strate-
environmental factors as a cue to offspring t- gies, he succumbed to subdivide heterochromatin
ness. In P. citri, several environmental factors into two types: constitutive heterochromatin
have been explored especially pertaining to (CH) and facultative heterochromatin (FH).
embryogeny and other biological features of As a constituent of chromosomal architecture,
female reproduction. These measures include the constitutive heterochromatin comprises consid-
role of population density (Varndell and Godfrey erable portions of the genome in higher eukary-
1996; Ross et al. 2010a, b), impact of tempera- otes that include specialized chromosomal
ture (James 1937; Nelson-Rees 1960), and age of domains that are endowed with repetitive DNA
3 Cytogenetics 37
sequence specicities (e.g., centromeres, telo- for the study of genomic imprinting, operating at
meres, and nucleolar organizer regions). CH at the level of a gene and/or at a single chromosome
the molecular level is marked by distinctive or a whole chromosome set, wherein inactivation
structural changes incurred at the level of DNA of (1) one of the two X chromosomes in female
sequences, and with active participation of con- mammals and (2) a male haploid set of mealybug
stituent histones, and consequently upon its chro- chromosomes in a complement, serve as a unique
matin remodeling. In addition, recruitment of example for the consideration of epigenetic
HP1 (heterochromatin protein 1) at times serves phenomena.
as an essential ingredient of heterochromatin There is good evidence that the control of
structure. The interactions and dimerization transcription involves active participation of var-
activities of HP1 with that of DNA sequences, ious proteins which bind specically to methyl-
RNA, and histone moieties using appropriate ated DNA, wring in histone modication
combinations bring about repressive chromo- complexes, and eventually in local chromatin
somal complexes. This situation is thought to be remodeling processes.
widespread in many eukaryotic genomes and in Considering these features, the differential
some instances, this appears to be a conserved chromatin formation during the course of chro-
genome component lending its role appropriately mosome inactivation processes in the case of
from yeast to mammals (Nokayama et al. 2001; mammalian females and in the case of the pater-
Nielsen et al. 2001). The CH could also be diag- nal chromosomal set in male mealybugs repre-
nosed by highly methylated DNA sequences, sents a very clear case and an outstanding genetic
and/or histone modications that are enriched manifestation offered in the studies pertaining to
with, for example, methylated lysines an effort to understand the modes and methodol-
(H3K9Me3) and yet in depleted form in the case ogies involved during such processes (FH
of both H3K4Me3 and acetylated H4 (acH4). formation).
As the name implies, facultative heterochro- This exceptional situation offers immense
matin comes into force or effectiveness upon academic help in eliciting more on these topics;
their need to undertake any exigency purposes Lakhotia (2004) made efforts to shortlist achieve-
(such as gene regulatory activities). FH is a ments dwelling upon ongoing excitements that
euchromatic component but upon developmental prevail in the arena of epigenetical phenomena
cues acquires a highly compacted chromatin contributing towards the current phase of knowl-
structure to transform itself into an heterochro- edge available regarding heterochromatinization
matic comportment. In its native state, FH is progression. Several recent reviews have been
devoid of repetitive DNA sequences. Facultative forwarded detailing the prospects of mechanisms
heterochromatin differs from constitutive hetero- and functioning of various epigenetical programs
chromatin with respect to DNA sequence rear- that incorporate during gene regulatory activities
rangements but not at the nucleosomal level. At (Surani 1991; Li 2002; Cairns 2007; Kouzarides
the nucleosomal level, FH has many molecular 2007; Skiniotis et al. 2007; Bell and Spector
features similar to CH. From the pointing of its 2011). Of particular signicant and recent prog-
impaction among higher organisms and in cyto- ress are achievements heralded in the case of epi-
genetic context, FH affects only one of two genetic regulatory activities during genomic
homologous loci or homologous chromosomes, imprinting programming of the mammalian
or homologous chromosomal set. X-chromosome (Sado et al. 2005). On the other
Genomic imprinting is dened as a parent-of- hand, the present review focuses on recent
origin specic expression of selected or affected achievements made in our current understanding
gene(s), and has generally been associated with of the role of DNA methylation, histone modi-
specic changes in DNA methylation proles cations, and some points upon the chromatin
and in histone modication processes. Even remodeling processes pertaining to genomic
though there are numerous examples available analysis (e.g., Planococcus citri /P. lilacinus)
38 R. Sompalaym et al.
have been amplied drastically due to spurious to control the whole of the inactivation program
developmental activities of insecticide (E4 & (Cattanach 1974; Lyon 1999; Brown et al. 1991).
FE4) resistance genes and thus, forcing upon Characteristically, the mammalian inactive
detoxifying esterases that have been spurred up X-chromosome shows a typical characteristic
due to spurt in DNA methylation processes (Hick organization as scored by micrococceal
et al. 1996; Field et al. 2004). Overt expression endonuclease treatment, because transcriptional
patterns of CpG methylation in these cases might factors do (or can) not bind to its condensed
have reected upon the amplication events of domains (Pfeifer and Riggs 1991). On the other
the concerned genes, the situation of such kind hand, chromosomes play a different role in view
may be considered reecting upon the mechanics of the situation that coccoid genomes have
of methylation processes associated with the offered as a readily packed and amenable mate-
copious presence of DNA transposons as was rial of chromosome research in any cytogenetic
found necessary in the cases of several verte- and/or biochemical exploration activities.
brates and in transgene experiments carried out One of the unique features while characteriz-
in plants (Feil and Khosla 1999; Field 2000). ing genomes is to introduce an enhancing mecha-
The historic ndings of Schrader (1921, nistic driving so as to yield differential
1923b) and Hughes-Schrader (1948) in which organization of homologous chromosomal sets
male chromosomes were found to be character- dwelling in one point of reference which allows
ized by the presence of a haploid chromosomal one to pursue gratuitously such as, for example,
set acquiring precocious condensation property to pursue more upon the mechanisms of sex-
and thus, becoming inactive ones (in Lecanoids) determination, genomic imprinting processes,
or put into an ordeal of genomic elimination (in and into inactivation progression (Hughes-
Diaspidids) during the course of embryogenesis. Schrader 1948; Chandra and Brown 1975;
Brown and Nelson-Rees (1961) described such Peterson and Sapienza 1993). For example, the
an event occurring by elaborating on chromo- mealybug genome is unique because it is in pos-
somal mechanics imposed upon heterochromatic session of unusual chromosomal characteristics,
components by means of undergoing a faculta- involving diffuse centromeric organization
tively heterochromatinization program. (holokinetic activity) that encompasses inverse
The condensation property of the paternal meiotic processes, leading to a signaling of an
chromosomal set of mealybug chromosomes is unorthodox mode of cell-cycle manipulation in
correlated in parallel with the expression for males (Hughes-Schrader and Ris 1941; Brown
maleness. In mealybugs and other coccoids, and Nur 1964; Nur 1990). Thus, some of these
radiation-induced chromosomal fragments are unusual genetic bounties could have driven
not lost during mitosis but persist as stable enti- Chandra and his collaborators in attempting and
ties in nuclei of both sexes, demonstrating that exploring further these genomic contents (e.g., P.
the centromere is diffuse and that freshly broken lilacinus or P. citri) at the DNA sequence level
chromosomal ends can still form telomeres or and of modied version of bases in the DNA
telomerelike structures and regulate associated sequence organization.
functions (Brown and Nelson-Rees 1961; Employing appropriate but standardized bio-
Chandra 1963a). When broken chromosomes chemical protocols (Jamaluddin et al. 1979;
were transmitted by fathers to their sons, each Achwal and Chandra 1982; Achwal et al. 1983,
chromosomal fragment underwent heterochro- 1984; Karnik 1983; Deobagkar et al. 1982, 1986)
matization progression suggesting the presence have enabled their fruitful extraction of total
of multiple centers of chromosome inactivation. nuclear DNA content based on an Indian
This situation contrasts with the condensation Planococcus genome. These assays were utilized
property exhibited in mammalian females, for the purposes of studying the primary nature of
wherein the inactive X-chromosome is identi- methylation status by means of HPLC and chro-
able with a single center of activity and is thought matography which enabled disclosing the pres-
40 R. Sompalaym et al.
ence of signicantly higher amounts of 5-methyl methylation processes and were found to occur
cytosines in some portions of the genome. This about 2.5 times more frequently than those show-
was veried by dinucleotide analysis in which ing female specic methylation sequences.
5-mC seemed over represented with respect to Bisulphite modied DNA samples revealed an
other sequences (viz., CpA, CpT, CpC). interspersion of CpG and non-CpG methylation
Unusually higher amounts of 6-mAdinosine among sex-specic methylated sequences. This
(6-mA), 7-mGuanosine (7-mG) were also study also pointed out that there were more non-
encountered (Deobagkar et al. 1982). Achwal CpG methylates and/or at least twice as many
et al. (1983) reported a new protocol to isolate sex-specic methylated sequences found in
and characterize antibodies raised specically to males than in females. They thus based on those
5-mC, 6-mA, and 7-mG, a situation rarely found sequences that there could be offering a closer
in higher eukaryotes at that time. With the use of association between sex-specic methylated
immunobiochemical approaches they were able sequences located in transcriptionally silent chro-
to evaluate the samples to the same level of con- matin zones and those assays resistant to DNase I
tention to that of higher eukaryotic samples (viz., zones.
Drosophila, Human, etc.). Scarbrough et al. (1984) studies were based
Devajyothi and Brahmachari (1989, 1992) on the differential levels of 5-mC in the males
present evidence of obtaining homogeneous and females of Pseudococcus calceolariae and P.
extraction of DNA-methyl transferase enzyme obscurus and thus they were able to relate their
that were found specic to the test material ndings and that these results driving them to
(Planococcus citri/ P.lilacinus). The enzyme arrive at conclusions that males display higher
extracts exhibited a proactive mode of action and incidences of methylated sequences than those of
found preference for salt extraction techniques, female samples. Kantheti (1994) describes, with
because that appeared equivalent to routine the help of specic antibodies raised against
extraction protocols utilized in the case of mam- 5-mC, that there were more 5-mC localization
malian methylase assays. These results demon- spots identiable on male cells than on female
strate that the enzyme assays have had high ones in the case of P. lilacinus. There were also
specicities for denatured DNA substrates. two more studies reported on Planococcus citri
Mohan et al. (2002) using random stretches of P. (Bongiorni et al. 1999; and Buglia et al. 1999)
lilacinus DNA sequences, the technique of which whose genomic exploration of P. citri samples
was found to be helpful in delineating repetitive related to the prevalence of sex-specic cytosine
sequence analyses that were inferred as higher specicities but arrived at conicting inferences.
than those of other conventional sequences and Khosla et al. (1996) present evidence suggest-
were also found much higher than those of ing existence of specic DNA fragments that
Drosophila samples scrutinized and compared were perhaps offering to serve as a primary sig-
wherein GCs were found less frequent. Thus, nal during the elaborate mechanism as a contrib-
they infer based on this situation that seemed uting factor towards chromosomal imprinting
promising for the considerations upon inuenc- activities. Chromatin organization of Planococcus
ing on CpG dinucleotide sequence frequencies lilacinus was chosen for the purpose of extrapo-
which was found exclusively in those genomic lating rather than to consider offering as contrib-
samples. Methylation specic arbitararily primed utory factors to their functional spectrum.
(MS-AP), polymerase chain reaction (PCR), and Digestion of P. lilacinus samples with micrococ-
subtraction hybridization protocols were found cal nucleases showed 35 % of the male genome
helpful to Mohan and Chandra (2005) and thus to samples were different and the same were
describe the isolation and sequencing of sex- assayed and found to be more resistant to the
specic CpG methylation sequences that were introduction of enzymatic activities; as such
prevalent in genomic DNA samples of P. lilaci- these samples were designated nuclease resistant
nus. These sequences showed male specic chromatins (NRCs) fractions. This component
3 Cytogenetics 41
was present invariably in both sexes and through- towards processing of the heterochromatization
out the genome. However, cloned NRC DNA program. It was also felt that some nrc51 frac-
contained A + T rich sequences that were found tions were not accessible to MNases even in
revealing some homology towards that of sam- euchromatic chromosomes. For the same they
ples of mouse - satellites. Salt fractionation offer the suggestion that these sequences might
techniques revealed that these sequences were have been inferred to contribute towards
found to be matrix-associated. Based on these centromeric-type activity; instead, they were
experiments, they were tempted to offer some found to be dispensed with all along the length of
solutions in the form of those DNA sequences the chromosomes. It was well known that a sin-
present explicitly in NRC fractions and it was gle inactivation center exists in the case of the
possible to infer that this sample would serve as a mammalian inactive X-chromosome, in contrast
resource material for a future course of genetic to the situation prevailing in the mealybug chro-
studies. Thus, Khosla et al. (1996) ndings thus mosomes exhibiting multiple centers along the
are directed towards offering these parameters length of individual chromosomes that serve as a
that could as well be serving as a mode of strategy model system for the chromosomal inactivation
and further to consider them as putative centers program. In the light of these ndings, these are
for initiation of facultative heterochromatization the distribution specicities for nrc50 and nrc-
processes. However, they also cautioned that there 51fractions over the mealybug chromosome sam-
are other contributory factors that they might ples and considering them for their presence in
interact with this grand executive operation. In the the form of several heterogeneous NRCDNA
meanwhile a thorough scrutinization is necessary fragments and of enrichment within the unusu-
and required in an extensive way prior to arriving ally organized chromatins of the male would
at any kind of generalization in this regard. raise the possibility of examining them and per-
With the help of southern hybridization and haps serving as putative nuclear sequence loci in
FISH techniques, Khosla et al. (1999) provide the form of expression of multiple inactivation
results proclaiming the extrapolation of NRCs centers.
and further about prevalence of subdivisions of Extending these experiments as an extrapola-
these fractions in the form of two middle repeti- tion undertaken by Khosla et al. (1999), they pro-
tive sequences, designated as nrc50 and nrc51 vide descriptions based on their explicit pattern
samples. It was also found that they were differ- of expression of this unusual chromatin organiza-
entially organized within NRC composition and tion designated as NRC fractions during the
more interestingly they have enabled distinguish- course of cytologically identiable regions and
ing the sexes based on the placement of differen- during spermatogenesis and especially over
tial proximity. The NRCs were also found sperm nuclei even though their expression was
resistant to both MNase and DNAase I treatment on a maternal background. Furthermore, it was
and thereby enable exhibiting indistinct patterns made possible for them to infer that this compo-
that may help in identifying two sexes. Their nent can perpetuate through mitotic and meiotic
enrichment in NRC accounted to contain 50 and progression.
83 % for nrc50 and nrc51 type, respectively. It also appeared interesting that differential
Thus, 2530 % of samples remain resistant in chromatin organization forms procured from the
males but none in females. It has been shown samples of the mealybug (Planococcus lilacinus)
consistently that NRC is associated with the provide an important biochemical tool in consid-
nuclear matrix. On a nuclear matrix isolation eration of assessing and identifying maleness or
platform regarding male and female sample femaleness based on the presence or absence of
nuclei, it was found evident that the NRC frac- NRCs from the total genomic organization. Thus,
tions were present only in males but not in based on this important biochemical discovery, it
females. They further imply that it is the pater- was made possible for Khosla et al. (2006) to
nally derived hypomethylation set that drives hypothesize and suggest a biochemical model
42 R. Sompalaym et al.
that may be able to answer some of the vexing 3.12.3 Molecular Cytogenetics
problems confronted by geneticists included dur-
ing the course of understanding genomic imprint- HP-1 (Heterochromatin Protein-1) is a nonhis-
ing mechanics. They are of the opinion that by tone chromosomal protein with two highly con-
regulating NRC as a discriminating organization served domains. The amino terminal
in the paternal/maternal genome, it becomes pos- chromodomain (CD) has the capacity to bind
sible to discriminate male- oriented cells from either mono-, di-, or tri-methylated histone moi-
those of females while attempting to recognize ety (e.g., lysines) of H3 or H4 or others. The car-
facultatively heterochromatinized chromatin boxy terminal chromoshadow (CS) domains
organization in one or the other sex. At this junc- are involved in mediating protein protein inter-
ture, their inference was to ascribe that in the pre- actions (Eissenberg and Elgin 2000; Lachner
ceding zygote formation, the zygote is in et al. 2001). Historically, HP-1 was identied and
possession of the paternal genome in the form of isolated originally based on Drosophila melano-
the NRC positive state and as such, the status of gaster polytene chromosome heterochromatin
heterochromatin is in the form of negative effect. regions and subsequently, were procured from
Subsequent to sixth cleavage divisions, the said several other sources and from several other
NRC-positive paternal genome acquires hetero- organisms, considering these format posed us as
chromatization status based on the developmen- the basis for isolation and they were acquainted
tal decision made at some point in the ooplasm, through to the cloning experiments. By raising
in order to acquire a decision either to procure or antibody (CIA 9) against those subdivisions of
lose heterochromatin mediating proteins, thereby several homologues were procured. HP-1 are
acquiring a specic functional role based on a highly conserved and play a role in gene silenc-
NRC-positive or negative fraction. ing efforts in a diverse range of organisms (Singh
Subsequently, Mathur et al. (2010) present a and Georgatos, 2002). There appear to have been
genomic organization of another pseudococcid, instances wherein euchromatic zones require
Maconellicoccus hirsutus, thereby evaluating the HP-1 s for stabilization of their elongating tran-
obvious presence of the effective NRC fraction scripts (Vakoc et al. 2005).
and its mode of association with that of nuclear Epstein et al. (1992) were keen on extrapolat-
histone matrix content. They insist based on pre- ing the molecular biology of HP-1 and their ef-
vious experience that the afnity patterns ciency towards cloning and thus isolated several
between NRC and histone matrix form an impor- patterns of expression from Drosophila HP-1
tant binding property for a meaningful differen- homologues and the same were used to compare
tial expression especially eliciting developmental with samples drawn from several other sources
courses promoting the paternal mode of inheri- wherein their genomes were known towards
tance. The exhaustive study revealed by means of exhibiting heterochromatin programs in which
extraction and the identication of H3K27Me3, the role of HP-1 takes dominance. Because they
H4K20Me3, and H3K9Me3 proteins in both in knew that the degree of similarity between chro-
male- and in female-based samples and with a modomains (of polycomb) and HP-1 at the
signicant enrichment of H3K27M3 in the nucleic acid level it was found sufcient to detect
nuclear matrix of males compared to that of and isolate other genes from other organisms
females form an important and critical contribu- using low-stringency nucleic acid hybridization
tion. This particular biochemical component (Singh et al. 1991). Epstein et al. (1992) were
seems pointing towards and directing a cell-based exploring the possibilities of procuring HP-1
signal for a male sex-specic discriminating fac- homologues from several other sources; however,
tor. Furthermore, the analysis of cytologically they preferred to examine HP-1 s from mealybug
sorted nuclei indicates the presence of NRC in genomes because it was well-known that these
nuclei with different DNA content including the scale insect provide a robust example for such kind
haploid nuclei from males, is another interesting of consideration and thus may serve as a suit-
phenomenon disclosed in this genome. able target (Hughes-Schrader 1948; Nur 1990).
3 Cytogenetics 43
Thus, the coccoid genetic system is well recog- chromatin suppression. An extension of such
nized as one of the rst examples to pursue for kind of exploration made on the mealybug
examining parent-of-origin (parental imprinting) genome (P. citri) was represented and shown by
specic effects; subsequently, other examples intense staining of DAPI; but male cells were
were perused for said purposes including humans highlighted by discrete staining localization
(Solter 1998). But Epstein et al. (1992) were able rather than that of interphase nuclei. Only ecks
to describe their attempts by means of molecular of stainability marks were found over the euchro-
characterization of two chromodomain- matic portions, but the representation at the male
containing proteins called PCHET-1 and prometaphase stage was by and large very clear
PCHET-2 (for putative coccid heterochromatin (Cowell et al. 2002). Thus, they made an asser-
proteins 1 and 2), from the mealybug genome, tion towards this effect that the role played by the
Planococcus citri. They were able to prepare HP-1 protein in silencing of concerned genes is
cDNA encoding these proteins realized in clon- thought to be a conserved function (Nokayama
ing and in which it was shown that PCHET-1 et al. 2001; Nielsen et al. 2001).
seemed to have more potential than that of Recent studies on methylated histones have
PCHET-2. This fusion product was later utilized revealed that the level of methylation of the spe-
for exploring the expression patterns of cic lysines may have an important functional
PCHET-1 in other mealybug tissues and it was consequence for the assembly of heterochroma-
conrmed that it assisted in a male tissue-specic tin formation. Acetylation and methylation are
manner. However,, the specicities of tissue dis- the two types of post-transcriptional modica-
tribution of this protein may suggest the most tions known that have been identied in histones
sought after gene, but it was not at the level of (Wu et al. 1986). The histone code is a sugges-
correlating to the extent of identifying the male- tion made in which covalent modications may
specic heterochromatic chromosomal set. be brought about by the kind and mode of the
Moreover, PCHET-1 was not found traceable on participation of chromosomal proteins and as
female cells. Thus, they opine that PCHET-1 in such, a modication will have effects on driving
combination with other factors may help in pro- towards tissue-specic expression patterns.
viding a role for the sex-determination device. Kourmouli et al. (2004) have made observations
Many decades of concentrated work on het- that on the N-terminal tails of lysine 20 of H4, it
erochromatization in terms of cytological and is trimethylation of this lysine that occurs; but if
molecular characterization reveal that this chro- it is dimethylation of lysines it was shown to be
mosomal component (whether constitutive or associated with euchromatic portions of the
facultative) consists based on a macromolecular genomes (Fang et al. 2002; Kourmouli et al.
mould in the form of a repressive chromatin com- 2004). Furthermore, Kourmouli et al. (2004)
plex (Spofford 1976). It is well known that meth- have reported that in the murine examples, the
ylation of lysine 9 of H3 by Suv (3)9 methyl trimethylated lysine 20 of H4 (but not the Me(2)
transferase creates a binding site for HP-1 (CD) K20H4) establish specic relationships in the
resulting in the formation of a repressive protein presence of Suv(3)9 histone methyl transferase
complex; since it was considered the most robust activity, with that of Me(3)K9H3 thereby
histone modications known. accounting for epigenetic crosstalk between H3
While attempting to elicit mutual relation- and H4. Extension of such kind of study revealed
ships existing between heterochromatin, HP-1, that in the coccoid examples analyzed as a target
and trimethylated lysine 9 of H3 (Me(3)K9H3) as for action it was expounded that its expressivity
a requirement in analyzing X-chromosome inac- was observed on the facultative heterochroma-
tivation program is resolvable us in the mamma- tized paternal chromosomal set. They made a
lian examples including humans, Cowell et al. detailed assessment of this situation by means of
(2002) observed that there were elevated levels DAPI stainings where the heterochromatic
of trimethylation at the notied sites resulting in component forms a brightly stained property
44 R. Sompalaym et al.
(Epstein et al. 1992; Bongiorni et al. 2001; geted for such an exploration in order to delineate
Kourmouli et al. 2004). In the female mealybug chromosomal differences, especially pointing out
cells, Me(3)K20H4 is found scattered uniformly DNA sequences pertaining to differences occur-
throughout the chromosomal set. ring at the organizational level, to the extent of
Most imprinted loci may have key regulatory identifying methylated and nonmethylated
elements that are methylated on one of the paren- chromosomes.
tal chromosomes. For several of these differen- Bongiorni et al. (1999) have made a detailed
tially methylated regions, recent studies establish account of the structural organization in respect
that the unmethylated chromosome has a special- to both males and females, and the paternal
ized chromatin organization that is characterized derived haploid set was found to be
by nuclease hypersensitivity. In such a situation, hypomehtylated to that of the maternally derived
the question is raised as to whether associated chromosome. In males it is the paternally derived
chromatin features regulate the allele specicity hypomethylated haploid set that is heterochroma-
of DNA methylation at those imprinting control tized. To their surprise, in female embryos, half
regions. of the chromosomal complement was under-
Taking cognizance of a lead from the bio- methylated and thus, they inferred that the under-
chemical front that was well demonstrated from methylated chromosomal set in females
the reports of Scarbrough et al. (1984) and represented was of paternal origin, emphasizing
Devajyothi and Brahmachari (1992) and its rele- that DNA methylation could be at the basis of
vance to the possibility of establishing preva- imprinting phenomena at the chromosomal level.
lence of relationships between two states, DNA Thus they suggest that the two haploid sets are
methylation processes and chromosome imprint- imprinted by parental-of-origin-specic DNA
ing phenomena in the coccoid genetic system is a methylation with no correlation with the known
jerk in our understanding of chromosome gene silencing properties of the base
imprinting phenomena and is considered monu- modication.
mental in coccoid genetic research. In order to In their next venture (Bongiorni et al. 2001),
probe further this important component of scale they carried out experiments based on western
insects, Prantera and his team (2012) have initi- blotting and immunolocalization with uorescent
ated unearthing several molecular cytogenetic microscope-level observations upon mealybug
complexities. Following is a descriptive account genome P. citri. Their intuition was to identify a
of their research accomplishments. cross-reactive protein epilope whose properties
In order to probe and enlighten based upon suggest that of a homologue of Drosophila HP-1,
implications of molecular and chromosomal present in this species. By analyzing the distribu-
level investigations undertaken by Bongiorni tion patterns upon immunouorescence spottings
et al. (1999) who made a beginning towards they could infer the distribution of this HP-1-like
prevalence of procuring knowledge of the P. citri protein in male and female cells during the cell
genome of Italian origin. They utilized the RE/ cycle and in the early embryogenesis. It was evi-
NT technique (restriction enzyme directed in situ dent to point out this (HP-1-like) protein colocal-
nick translation) upon exploring of DNA izes with male-specic heterochromatin, thereby
sequence-level organization, thereby extrapolat- implying that this protein plays a role in the pro-
ing the P. citri chromosome (Ferraro et al. 2001). cess of facultative heterochromatization.
Concentrating specically based on MSPI and its However, they allay some doubts as to the
methyl-sensitive isoschizomer HPa II when used nature of the presence of P. citri HP-1-like pro-
as nicking agents, led them to make incisions into tein in embryos of both sexes which had led them
the genome by exposing organizational differ- to infer a protein factor was involved in the rec-
ences prevailing between homologous chromo- ognition of the imprint signal, suggesting that at
somes and subchromosomal regions (Prantera least there could be another factor provision
and Ferraro 1990). The P. citri genome was tar- which was found to be involved in the induction
3 Cytogenetics 45
reversal occurs, the HP-2-like epitope is lost by lagging, abnormal condensation, segregation
the chromocenter remnants before the HP-1-like, defects), more so on structural maintenance com-
which thus seems to be insufcient to anchor ponents (SMCs).
HP-2-like to chromatin. It has also become evi-
dent that the strict colocalization of HP-2-like
with the chromocenter is not abolished in HP-1- 3.12.4 Chromatin Remodeling
like knockout embryos.
Molecular results based on some mammalian In many diverse organisms, gamete formation
examples, also including the mealybug genome, originates in a cytoplasmic, but highly conserved
were obtained independently by Kourmouli et al. structure, known as germ-line cysts. Germ-line
(2004) and Schotta et al. (2004) in an experiment cysts (or saclike structures) are composed of a
to certify the effect that Me(3)K9H3 employing group of cells; it is apparent that they took their
Me(3)K20H4 through the participation of HP-1 initiation from a single cell that underwent syn-
promoting heterochromatin formation appears to chronous cell divisions followed by incomplete
be a global-level event. But what was not clear cytokinesis. Modication of the chromatin struc-
about this was how HP-1 modulation is involved ture is one of the main epigenetic regulations con-
during gene activation processes in the case of ceived to carry out its operation in order to
the mealybug genome (P. citri; Bongiorni and undertake unique gene expression modalities. The
Prantera 2003; Bongiorni et al. 2007; Kourmouli male germ-line cyst is the organ that facilitates
et al. 2004). In contrast, acetylation of histone H4 executing the meiotic and/or post-meiotic mode
(AcH4) was found to be absent on the male- of gene regulation activity sharing during gameto-
specic heterochromatization processes (Ferraro genesis. The germ-line cyst morphogenesis
et al. 2001), whereas the depleted level of activa- acquires the responsibility of delivering the
tion of AcH4 was observed in the case of human respective genomic content to their destined sites.
X-chromosome inactivation (Jeppesen and Male meiosis of scale insects is interesting
Turner 1993). The foregoing issues have driven because meiotic sequence progressions proceed
to an understanding with a suggestion that Me(3) in accordance with those of inverse meiosis.
K9H3 via HP-1 to the Me(3)K20H4 pathway in Thus, during male meiosis each spermatogonial
an evolutionarily conserved mechanism of action precursor cell nucleus produces a bunch of syn-
for an epigenetic route to silencing large chromo- chronously dividing spermatogonia in a cytoplas-
somal domains by facultative heterochromatiza- mic cyst. Each spermatogonium divides four
tion (Chadwick and Willard 2004). times to produce a cyst of 16 primary spermato-
While establishing the prevalence of Me(3) cytes which then undergo two meiotic divisions.
K9H3 to HP-1to Me(3)K20H4 relationships in the Subsequently, each spermatogonium undergoes
case of P. citri genomes, Bongiorni et al. (2007) the rst equational and then the second reduction
proceeded further to interrelate the position of the division, which is characterized by specialized
HP-2-like protein (PCHET-2) based on RNAi movements directed and dictated by some
experiments. With the intermediation of ds RNAi unknown sources. But recent studies undertaken
(Fire et al. 1998) and by interference of knocking by Buglia and Ferraro (2004), Buglia et al.
down PCHET-2 in P. citri embryos, it was resolved (2009), and Bongiorni et al. (2009) have pro-
that the consequential depletion of the heterochro- vided some clues to learn more about the extent
matization pathway resulted in deheterochromati- and nature of expression, wherein these chromo-
zation with respect to gut cells and Malpighian somal movements were maintained and manipu-
tubules, whereas Hp-1 and Me(3)K20H4 in the lated by the monopolar spindle in which
same nuclei are either dispersed or absent. microtubules make physical connection with the
Embryos treated with ds RNAi (double-stranded euchromosomal set, rather than with the hetero-
RNA interference) targeting PCHET-2 also exhibit chromatic component as was contended earlier
chromosomal abnormalities (such as chromosome by Hughes-Schrader (1948). Even though Sciara
3 Cytogenetics 47
chromosomes practice monopolar spindle activi- H4K20Me3. This was found to be a consistent
ties, it seemed to be maintained through the expression pattern until the spermatid formation,
occurrence of monokinetic activity wherein mei- thereby demonstrating the supremacy of histone
otic products were manicured by sister-chromatid modications throughout the male part of the
cohesion (Esteban et al. 1997). meiosis. This situation is in congruence with that
By utilizing antibody-specic tracings, Buglia of Khosla et al. (1999, 2006) observations and of
and Ferraro (2004) describe an immunouores- their proposals advocating the presence of NRCs
cent staining protocol backed by enhanced active on paternal cell lineage until sperm maturation.
participation of fusomal elements, such as By now, it seems evident by pointing out that by
F-actin, included in the elaborate descriptions of the end of spermatogenesis PCHET-2 may be
factors that demarcate local morphogenesis losing its grip. Bongiorni et al. (2009) contend
essentially demarcating the cytoplasmic compo- that the presence and supremacy of H3K9Me2 &
sition of the male germ-line cysts. The co- 3 methylation processes dominate throughout the
localization of all these factors is an indication of course of gametogenesis, and with respect to the
the triggering action that could be measured by content of these proteins, they are in disagree-
densitometric proles which further enable in ment with the contention of Buglia and Ferraros
providing descriptions about the prevalence of (2004) observations. This pertains to the quan-
two kinds of sperms emerging but equipped with tum of differential distributions regarding
variable loads with respect to individual sperm euchromatic spermatids, because these products
content. take their origin from a single meiotic event.
A continuing search by Bongiorni et al. (2009) However, Buglia and Ferraro (2004) strongly
proceeded towards extrapolating procurement of dene that values they procured were essentially
the resources to be used during the gametoge- based on densitometric tracings, citing differing
netic processes and seemed to be in possession values with respect to H3K9Me2 & 3 and of
until early embryonic development. CIA9.
Immunolabeling of such components in order to In their subsequent study, Buglia et al. (2009)
probe has enabled identifying the presence of have elaborated mustering of resources pertain-
protein components such as H3K9Me2 & 3, ing to the development of female phases of
H4K20Me3, HP-2, and PCHET-2-like, that were gametogenesis of P. citri. Their results provide
concentrated in the paternal part of the meiotic the presence of a proteic component; this time the
stages throughout, but not in the female line to presence of HP1 and Su (var) 39 (a different
the extent of oocyte formation. On the other chromosomal protein), makes all the more impor-
hand, there were no traces of these modiers in tant contributions occurring during female gam-
the female gametogenesis. The redistribution of ete formation. Pertaining to the deposition of
epigenetic signaling marks in spermatids might variable contents of eggs it was found to contain
be related in the tracings of the processes con- two different kinds of cell inclusions, deposited
cerned with the establishment of parental imprint- in eggs, thereby categorizing in such a way as to
ing. Bongiorni et al. (2009) narrate the modes of act differently upon different ages of females.
operation through to the entry of sperm into the Based on these biochemical characteristics,
oocyte environment, where they are in posses- females with 40-days older age were considered
sion of distinct H3K9Me2 and 3 methylation as a younger group and those of 80-days old as an
marks that were found in the early pronucleus. older (aged) group. The ndings of larger
Observations were made of such kind of effect amounts of epigenetic factors accumulated in the
during the course of spermatogenesis indicating group of aged females in comparison to the
the presence in the form of heterochromatic com- younger ones was found to be an important
ponents decorated by H3K9Me2 & 3 and deciding factor. These studies have led to the
PCHET-2. Regarding the euchromatic compo- supposition of playing as a primary role based on
nent, it was shown containing HP-2-like and differential maternal contribution.
48 R. Sompalaym et al.
It was observed that the concept of genomic lated that the role of the mothers cytoplasmic
imprinting phenomena seemed to be lending environment might have been inicted by impo-
effective support in the cases of both Sciara and sition of environmental disturbances. Thus, dur-
coccoids, that the primary sex determination ing routine life-style courses, a one-to-one ratio
mechanism relied upon considering conse- in the case of sexually reproducing and one-to-
quences of occurrence of chromosome imprint- none in the case of parthenogenetic system, the
ing (Chandra and Brown 1975; Brown and sex ratio will operate in an expected line.
Chandra 1977). It also appears obvious in the However, if any change is incurred with respect
case of mealybugs, that possibly it was at the to the sex ratio it could possibly be envisioned
instance of mothers that enable directing and dis- and perceived as operating under constraints due
criminating the sex of her offspring. In addition, to the external forces that thrust upon maternal
the extension of this provision should yield environmental cues.
mechanistic support to the concept that mater- Currently, the mechanism of the genomic
nally controlled sex determination could also imprinting phenomenon is still unclear although
give way or leverage for control of the progeny the role of PCHET-2 and histone modications
sex ratio. Earlier, Nelson-Rees (1960) had con- seems evidently involved in effecting the faculta-
tended that the sex ratio in mealybugs uctuates tive heterochromatization process in the case of
among females and is markedly inuenced by the mealybug genome (P. citri). Females may
mothers age at conception towards the brood. In volunteer and might offer to alter the concentra-
both sexually reproducing and in parthenogenetic tion of those proteins in their eggs to their con-
mode of reproduction, the imprinting process ini- tention so as to modulate the sex ratio of their
tiates at and in the egg cytoplasm at the time of broods. Along this line, Buglia and Ferraro
the fertilization program. (2004) and Buglia et al. (2009) observations
Parental genomic inuences on the fate of off- point towards the situation that under the varied
spring development are evident in both inverte- concentration of CIA9-based positively stained
brate and vertebrates. Maternal effects are protein and of those of observations pertaining to
commonly mediated through deposition of the the eggs of females possessing variable amounts
cytoplasmic transcripts essaying protein products of proteins and at variable ages prior to mating
in oocytes during oogenesis in the female germ- should bring forth more differences in the egg
line. These then exert their effects on the fertil- chamber. They also apprise that females would
ized eggs and drive impulses upon early produce male-biased offspring whereas the oppo-
embryonic developmental processes (De Robertis site effect of maternal aging prior to mating was
et al. 2000; Gosden 2002), unlike some mamma- also observed in other studies.
lian examples that may provide guidelines for Prantera and Bongiorni (2012) postulated that
any kind of eventuality (Li et al. 2008). However, the embryonic cytoplasm at the blastoderm stage
there are no specic studies undertaken pertain- determines whether the paternal chromosomes,
ing to the operating mechanisms responsible for which are marked by DNA hypomethylation and
maintenance of genomic methylation imprints, H3K9me3 methylation marks, could be able to
even though the P. citri genome may serve as drive towards undergoing heterochromatization
very good material for such kind of expeditions. processes or not, and thereby giving rise to a
In view of this trepidation, it is possible to male or female embryo, respectively. Given the
infer that the mother can embark upon an initia- causative role and presence of PCHET-2 on
tion or directing a particular path towards the male-specic heterochromatin formation and
choice of her offspring and its bestowing effec- also based on the amount of PCHET-2 in the
tiveness on the sex-ratio potential. In promulgat- developing embryo, may prefer it as a crucial
ing the imprinting phenomenon, in terms of factor to drive the embryo either towards male-
evolutionary consequences with reference to a ness or femaleness. It is already envisioned that
choice-based progeny sex ratio, it was also postu- the effectiveness of facultative heterochromatini-
3 Cytogenetics 49
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Taxonomy
4
M. Mani
The insects coming under Hemiptera, Phenacoccinae, and Rhizoecinae (Downie and
Sternorrhyncha, Coccoidea, Pseudococcidae, Gullan 2005). This estimate was recently revised
and Putoidae are named as mealybugs (Williams in light of integrated molecular and morphologi-
2004). Following the application by Miller cal data, and only two subfamilies emerged:
(1975b), the family-group name Pseudococcidae Pseudococcinae and Phenacoccinae (Hardy et al.
(Cockerell 1905) was placed on the Ofcial List 2008). Molecular studies may either verify this
of Family-Group Names in Zoology (Melville grouping or show a different picture. By the study
1983) (type genus Pseudococcus Westwood of prokaryotic primary endosymbiont
1840). A (1968) attempted a higher classica- (P-endosymbiont) nucleotide sequences, Thao
tion of the Pseudococcidae based on a study of et al. (2002) showed that Antonina pretiosa
the characters of adult males of 17 species. The Ferris, presently included in the Sphaerococcinae,
scale insects are generally divided into two is closely related to the blue-green or blue-black
groups, namely the archeococcids and the neo- mealybugs of the genera Amonostherium
coccids. The archeococcids possess two to eight Morrison and Morrison, Australicoccus Williams,
pairs of abdominal spiracles, which are absent in Melanococcus Williams, and Nipaecoccus Sulc.
the neococcids (Koteja 2008). The family These genera are included in the Trabutininae, as
Pseudococcidae (mealybugs) belongs to the neo- discussed by Koteja (1974a, b).
coccid group. Pseudococcidae constitutes the second largest
After extensive studies on the labium of 84 family of Coccoidea, with more than 2000
species of Pseudococcidae, Koteja (1974a, b) described species and ca. 290 genera (Ben-Dov
proposed that the family is composed of four sub- 2006; Downie and Gullan 2004). Pseudococcids
families: Trabutininae, Rhizoecinae, occur in all zoogeographical regions of the world.
Sphaerococcinae, and Pseudococcinae. This Pseudococcids are distributed in different geo-
classication has gained wide acceptance. A graphical regions as follows: Australasian region
recent phylogenetic study, based on the analysis (459 spp.), Afrotropical region (298 spp.),
of nucleotide sequence data, supported the exis- Nearctic region (424 spp.), Neotropical region
tence of three subfamilies: Pseudococcinae, (283 spp.), Oriental region (431 spp.), and
Palearctic region (710 spp.). Of the described
species, pseudococcids are most abundant in the
M. Mani (*) Palearctic region and least numerous in the
Indian Institute of Horticultural Research, Neotropical area. There are about 2000 species of
Bangalore 560089, India mealybugs worldwide. In southern Asia, 353
e-mail: mmani1949@yahoo.co.in
species of mealybugs have been recorded under considerable taxonomic value. This is exempli-
61 genera of which 105 species occurred in India ed in certain species of Chorizococcus and
with maximum species reported from Karnataka Spilococcus. The distance between the base of
(40 species) followed by Tamil Nadu (35 spe- one antenna and that of the other is of consider-
cies). India is rich in species of Formicococcus able value in separating species, particularly in
Takahashi (10 spp.), Antonina Signoret (six spp.), certain members of Rhizoecus. Most coccidolo-
Dysmicoccus Ferris (12 spp.), and Paracoccus gists have placed little taxonomic emphasis on
Ezzat and Mc Connell (six spp.). The genera, the eyes of mealybugs. However, the presence or
which are endemic to the Indian region, are absence of eyes in Rhizoecus and related genera
Aemulantonina Williams, Coccidohystrix has proved to be taxonomically useful. The
Lindinger, Eriodes Green, Lankacoccus labium varies considerably in shape and form
Williams, Pedronia Green, and Pseudantonina and may be elongate and slender in some species,
Green (Williams 2004). while in others, it is short and broad; in certain
Identications of mealybugs in practically all species, there appear to be signicant differences
cases are based upon the adult female. A sum of in the shape of both the basal segment and the tip
the morphological characteristics of the mealy- of the rostrum. Some species exhibit a sclerotized
bug used in identication of species belonging to area on the derm just anterior to the clypeus. In
family Pseudococcidae are as follows: Anal ring some instances, this has been of taxonomic assis-
is always present, divided longitudinally into two tance. The body form normally elongate; legs are
halves, each with single inner and outer rows of normally present and usually well developed.
angular cells and three setae; in a few instances, Considerable taxonomic emphasis has been
the ring is very much reduced, the sclerotization placed on the mealybug legs in the past by certain
is slight, and the pores apparently are nearly or coccidologists. Denticle or tooth on the plantar
completely lacking. Two pairs of dorsal ostioles surface of the claw offers an especially excellent
are normally present in the adult female. In some key character for the recognition of this series of
species, the number of these structures may be genera. Although the denticle or tooth still gener-
reduced, or they may be entirely lacking; thus, in ally is quite helpful in dening the members of the
some forms, the posterior pair is clearly present, genus Phenacoccus, it cannot be completely relied
but the anterior pair is lacking; in a few forms, upon as exemplifying this group alone. In many
the ostioles seem to be lacking in the adult but are species of Chorizococcus and Spilococcus, this
present in the rst stage. Others lack ostioles in tiny denticle or tooth on the claw is present, and
all stages; nevertheless, the totality of their char- it occurs in combination with other characters
acters places them in this family. The antennae that are not at all typical of the Phenacoccus series.
have been used in diagnoses of mealybug genera The body is normally with lateral groups of
for a long time and even for species separation. pores and has enlarged, conical setae, which form
Hence, more emphasis has been placed on the cerarii, that at times are evident only on the anal
antennal structure than on any other physical lobes, occasionally lacking, normally with pores
detail. In Antonina and related genera and species of the trilocular type present, rarely lacking.
of Eumyrmococcus, antennae may be reduced to Tubular ducts of a distinctive type are normally
one or two segmented stubs. It has been noted present as cylindrical invaginations in the derm,
that in certain genera, exclusive of Rhizoecus, the tube usually more heavily sclerotized at its
Geococcus, and Pygmaeococcus, where the opening, and with one side of the inner end of
antennae are noticeably short, small, ve to six tube showing a delicate lamentous prolonga-
segmented, and geniculate, the comparative slen- tion. Combinations of these characters will dene
derness or stoutness of the normal cylindrical the few aberrant forms of mealybugs that are
antenna in relation to its length has proved of known to exist.
4 Taxonomy 57
Translucent dots or pores on the hind femur 1. Place the entire specimen in a 6-ml., 1-in.
and tibia have denite signicance for species handled porcelain casserole dish approxi-
segregation. In some mealybug forms, each hind mately half-lled with Essigs Aphid Fluid
coxa bears a cluster of pores at its base, and the (see formula below), cover with 1-inch
area in which these occur is usually wrinkled. watch glass, and heat (120130 F) to dis-
This pore cluster is taxonomically important for solve (1015 min). A lateral incision made
differentiating certain species. The trochanter between the mid- and forelegs will help to
usually possesses a long seta at inner distal end clear the specimens more rapidly.
and the variation in its length and thickness 2. Remove the porcelain dish from the hot plate
proved to be a useful distinguishing character in and tease out the body contents while the uid
species of the Allomyrmococcini. The stoutness is still hot.
or slenderness of the pseudococcid legs in rela- 3. Transfer the cleared specimen to a droplet of
tion to their length has proved to be of much gum-chloral hydrate or chloral-hydrate
taxonomic importance. medium (see formula below). Apply a cover
Clypeolabral shield structure seems to reach slip and heat the slide on hot plate until the
its greatest development in certain bamboo- medium boils slightly. The specimen is then
feeding species of the tribe Serrolecaniini, and conditioned for examination under the com-
sometimes reaches almost the same length as the pound microscope. (Polyvinyl alcohol is also
clypeolabral shield. The structure is now known considered a good temporary-type medium.
to occur in many species but sometimes it is The specimens should be transferred from
barely perceptible. The extension is present Essigs Aphid Fluid directly into the solution.)
mainly in grass-infesting species and occasion- Valuable specimens are recoverable from this
ally in mealybugs feeding on the other groups of medium for permanent embedding in Canada
monocotyledons but, apparently, never in balsam, although this should not be delayed
dicotyledon-infesting species. longer than 3 or 4 months.
Cerarii situated on the dorsum of the body,
their total number, the number of enlarged coni-
cal setae, the presence or absence of auxiliary
setae, and the presence or absence of the accom- 4.2 Permanent Mounts
panying sclerotization have proved to be impor-
tant specic taxonomic characters. The following steps are used in the preparation of
Ventrally, the anal lobe is often sclerotized, permanent pseudococcid mounts:
and the character of this pigmentation is some-
times used as a taxonomic feature at the generic 1. Place the entire specimen in a 6-ml., 1-in.
and specic levels. handled porcelain casserole dish approxi-
The presence or absence of a circulus is mately half-lled with Essigs Aphid Fluid
exceedingly helpful as a key character within a (see formula below), cover with 1-inch
genus. At times, they vary in size, form, or num- watch glass, and heat (120130 F.) on a hot
ber to such a degree as to be of taxonomic value. plate until the body contents are dissolved
In some genera, several circuli may be present. (30 min to 1 h). A lateral incision made
between mid- and forelegs will help to clear
specimens more rapidly.
4.1 Temporary Mounts 2. Remove the porcelain dish from the hot plate
and tease the body contents while the uid is
The following steps are used in the preparation of still hot. If the specimen is not thoroughly
temporary pseudococcid mounts: cleared, tease out the loosened body contents
58 M. Mani
and transfer the specimen to a fresh solution of 6. Transfer the specimens to cellosolve in a clean
Essigs Aphid Fluid. Add three or four drops of depression slide and leave in this solution for
the prepared staining solution, consisting of not less than 5, preferably 20, min.
either acid fuchsin, lignin pink, or erythrosine 7. Transfer the specimens to xylene in a clean
(stain No. 2) (see staining solution formulae depression slide and wash thoroughly for 1 or
below), or even more of this solution if a deeper 2 min.
staining is desired. Heat again until the speci- 8. Place the specimens in a droplet of Canada
mens have absorbed the stain (1530 min balsam on a glass-cover slide and apply the
depending on the specimens involved.) cover slip. Use as little balsam as possible to
3. Transfer the specimens directly into a clear facilitate examination under the compound
6-ml porcelain casserole dish half-lled with microscope, especially under the oil-
tetrahydrofuran (C4H8O) and tease out the immersion magnication.
remaining body contents and excess stain.
Leave the specimens in tetrahydrofuran for The formula used to prepare Essigs Aphid
not more than 5 min, since prolonged periods Fluid is as follows:
may cause shriveling. In the case of very frag-
ile specimens, add a few drops of tetrahydro- Lactic acid (reagent grade 85 %) 20 parts
furan to the Essigs Aphid Fluid before Phenol (saturated in distilled H2O) 2 parts
transferring straight to tetrahydrofuran; this Glacial acetic acid 4 parts
will prevent shriveling. Water (distilled) 1 part
4. Transfer directly into Canada balsam and
apply the cover slip. Specimens should be The formula used to prepare chloral-hydrate
transferred rapidly form tetrahydrofuran to medium is as follows:
balsam, as little carry-over of tetrahydrofuran
as possible. Air bubbles are often left under Gum arabic 1g
the cover glass because the solution evapo- Dextrose 1g
rates quickly, but they will ultimately work Chloral hydrate 10 g
their way to the edge of the cover glass. It is
advisable to burst these bubbles with a needle Iodine crystals 1/10 g
dipped in tetrahydrofuran solution before Glycerin 1 cc.
placing the mount on heat to cure. This curing Water (distilled) 1 cc.
should not be done at more than 100 F, and
30 min to 1 his required to sufciently harden The formulae used in preparing the staining
the mount. solutions Nos. 1 and 2 are as follows:
5. When the balsam has hardened, the cover slip
may be ringed with shellac or other suitable No. 1 Essigs Aphid 15 ml.
Fluid
media to prevent later fracturing of the bal-
Acid fuchsin (2 % aqueous 20 drops
sam. It is good to remember that the clearing solution)
and straining process cannot be hurried. No. 2 Essigs Aphid 15 ml.
However, when the specimens are properly Fluid
cleared and stained and the mounting tech- Acid fuchsin (2 % aqueous 20 drops
niques are mastered, excellent mounts will solution)
result. Because of the high volatility of tetra- Lignin pink (2 % aqueous 20 drops
solution)
hydrofuran, some of the smaller and more
Erythrosin (2 % aqueous 20 drops
delicate mealybugs tend to collapse when
solution)
transferred into it. In such a case, step 3 should
be modied as follows:
4 Taxonomy 59
Staining solution No. 1 gives excellent results. the Australian mealybugs by Williams (1985),
One slight drawback, however, is that the speci- and the mealybugs of Central and South America
mens from certain lots may begin to fade after 3 by Williams et al. (1992). A systematic catalog of
or 4 months. Preliminary observations made over the mealybugs of the world (Insecta: Homoptera:
approximately a 2-year period indicate that the Coccoidea: Pseudococcidae and Putoidae) has
staining solution No. 2 tends to overcome this data on geographical distribution, host plants,
feature, at least to some degree. It is interesting to biology, and economic importance by Ben-Dov
note that the species vary in their response to (1994) and the mealybugs of Southern Asia by
staining, some turning to darker red than others Williams (2004). The above keys may be referred
after the same time in the staining solution. This for identication up to species level.
has been advantageous in certain instances, espe-
cially where two species are mixed on a single
host and are indistinguishable from each other 4.3 Field Identication of Major
when collected in the eld. In such instances, the Species of Mealybugs
specimens may be easily segregated by species
before they are mounted. All mealybug species resemble each other to the
Brief instructions for slide-mounting scales untrained eye, so it is very important that an
and mealybugs have also been provided by the expert is brought to identify the mealybug spe-
United States Department of Agriculture (USDA) cies involved. The following characteristics are
in their Systematic Entomology webpage. Keys useful for eld identication. The adult female
available in the identication of mealybugs are mealybug is considered for the identication of
the mealybugs of California by McKenzie (1967), body shape, size, and color (Table 4.1):
Table 4.1 List of mealybug species with eld-identifying characters with their respective images
Mealybug species Field characters Images of mealybug
Antonina graminis (Maskell) (Rhodesgrass mealybug)
Broadly oval to circular body; rotund in lateral view; dark
purple or brown body; without lateral wax laments;
enclosed in a white, felted sac that turns yellow with age;
usually with a long, slender, white waxy tube protruding
through a hole in the ovisac at the posterior end of the body.
Usually present on the crown or nodes of the grass host.
Ovoviviparous; rst instars are cream colored; legs absent.
(continued)
60 M. Mani
(continued)
4 Taxonomy 61
(continued)
62 M. Mani
Palmicultor browni
Body reddish brown to pink; about 3 mm in length; with
side (lateral) wax laments; no ovisac produced.
(continued)
4 Taxonomy 63
(continued)
64 M. Mani
(continued)
4 Taxonomy 65
(continued)
66 M. Mani
(continued)
4 Taxonomy 67
(continued)
68 M. Mani
Rastrococcus invadens
Ovoviparous; a tuft of hairs in the anterior region; lateral
laments increase in length from anterior to posterior region;
infestation conned to midrib of the leaves.
(continued)
4 Taxonomy 69
(continued)
70 M. Mani
Stemmatomerinx acircula
Body gray with white wax; about 23 mm long; some wax
seems to be lamentous; no lateral wax laments produced.
The number of wax laments protruding from The second type is root mealybugs/soil mealy-
the side of the body. bugs/subterranean mealybugs living in the soil
Presence and length of wax laments at the and feeding on the roots.
end of the body (i.e., terminal wax laments).
Color of eggs (if present).
Presence of an ovisac (a waxy mass covering 4.4 Role of Taxonomy
the eggs). in Management
Stripes on the body. of Mealybugs
Color of uids when crushed.
Success in pest management tactics including the
There are two types of mealybugs. One is leaf biological control programs depends on the cor-
mealybugs/foliar mealybugs/arboreal mealybugs rect identication of both the biological control
infesting the plant parts above the ground level. agent and the pest species. In last few decades,
4 Taxonomy 71
there have been more than ve major outbreaks Acerophagus coccois Smith (Bento et al. 1999).
of mealybugs causing alarming damage to crops, The most trenchant point concerning the parthe-
as a result of accidental introduction. nogenetic species P. manihoti is that an outbreak
The pink cassava mealybug, Phenacoccus could occur in southern Asia with the accidental
manihoti Matile-Ferrero, appeared on cassava in introduction of just a single immature specimen.
Africa in 1973 and soon spread throughout the Following the introduction of the cassava mealy-
whole cassava belt. To locate the source of this bug into Africa, another introduced mealybug
mealybug and to compare it with a morphologi- appeared in West Africa in 19811982, causing
cally similar species, which causes almost identi- extensive damage to fruit trees including mango.
cal damage to cassava in northern Brazil and This mealybug was initially identied as an
Guyana, required a considerable amount of time. undescribed species already known from India
Specimens from Africa and northern South and Pakistan and was later described as
America on the microscope slides showed wide Rastrococcus invadens Williams (Williams 1986).
variation in characters, and the method adopted This species is usually scarce in some parts of
to ascertain any limits to this variation required India because it is controlled by the natural ene-
rearing cultures in the laboratory at different tem- mies (Narasimham and Chako 1988); the intro-
peratures. This method normally induces wide duction of the encyrtid Gyranusoidea tebyi Noyes
morphological variation in mealybugs, helping to from India to West Africa and its swift control of
determine the limits of environmentally induced the mealybug are hailed as another biological
variation (Cox 1982; Cox and Williams 1981). control success (Neuenschwander et al. 1994).
The knowledge that the species in Africa was Another mealybug species was introduced
pink-bodied and uniparental and the species from accidentally to the Caribbean area in 199394
northern Brazil and Guyana was yellow-bodied and has since then spread beyond, eventually
and biparental (later described as Phenacoccus reaching USA. This damaging species was rap-
herreni Cox and Williams) was obscured initially idly identied by taxonomists as Maconellicoccus
because the specimens used for this study were hirsutus (Green); its biological control was
dead and had been preserved in spirit, obscuring described in detail by Kairo et al. (2000), with
the body color. When the two species could be discussion of the costs and benets. M. hirsutus
identied satisfactorily on the microscope slides, is widely distributed throughout the southern
it became apparent that the pink cassava mealy- Asia, Africa, and other parts of the Old World
bug, P. manihoti, was present in Paraguay and including Australia, and is still causing damage
Bolivia (Williams et al. 1981); hence, a search for in some parts of India. The introduced natural
natural enemies could be implemented there. The enemies, mainly the parasitoids Anagyrus kamali
introduction of the parasitoid Apoanagyrus lopezi Moursi (already known in the Old World) and
(De Santis) from South America to Africa and the Gyranusoidea indica Shafee, Alam and Agarwal
success of the biological control program against (collected in Egypt), and the predator
P. manihoti were well documented by Cryptolaemus montrouzieri Mulsant, have
Neuenschwander and Herren (1988) and Herren brought the mealybug under control. In response
and Neuenschwander (1991). Thus, the taxo- to this outbreak, an identication manual for the
nomic information can be retrieved in case the area (Watson & Chandler 1999) and a taxonomic
mealybug introductions originate from this area. study of all the instars of M. hirsutus (Miller
Phenacoccus manihoti remains a threat to the 2002) were produced.
cassava areas of southern Asia, as does the yellow Yet another mealybug is causing concern in
cassava mealybug, P. herreni, which still causes the Caribbean area Paracoccus marginatus
problems in South America. Reduction of P. her- Williams and Granara de Willink, described from
reni populations is now under way, mainly Mexico and parts of Central America as recently
through the introduction of the parasitoids as 1992, has become a serious pest in the
Apoanagyrus diversicomis (Howard) and Caribbean islands, where it attacks numerous
72 M. Mani
Table 4.2 List of mealybug species, correct identity of which led to a successful biological control
Country of accidental Introduced parasitoid for classical biological
Mealybug species introduction control
Phenacoccus manihoti Matile-Ferrero Africa Apoanagyrus lopezi (De Santis)
Phenacoccus herreni Cox and South America Apoanagyrus diversicornis (Howard)
Williams
Rastrococcus invadens Williams West Africa Gyranusoidea tebyi Noyes
Maconellicoccus hirsutus Green USA Anagyrus kamali Moursi
Paracoccus marginatus Williams and Caribbean islands Acerophagus papayae Noyes and Schauff
Granara de Willink Pseudleptomastix mexicana Noyes and Schauff
Anagyrus loecki Noyes and Menezes
Paracoccus marginatus Williams and India Acerophagus papayae Noyes and Schauff
Granara de Willink Pseudleptomastix mexicana Noyes and Schauff
Anagyrus loecki Noyes and Menezes
plant species, especially papaya (Carica papaya). come under the existing keys, it may be named as
The mealybug has now reached the southern a new species.
USA. A search for natural enemies in Mexico
(Becker 2000) located three parasitoids that are Acknowledgement Dr. Sunil Joshi, Principal Scientist,
now in use in controlling the mealybug. An off- NBAIR, Bangalore is acknowledged for providing
information.
shoot of the biological control program has been
a detailed study of all the instars of P. marginatus
by Miller et al. (2005). The mealybug had
References
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Entomol Suppl 13:1210
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staining and permanently mounting small arthropods. Natural History Museum/Southdene Sdn. Bhd,
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mealybug and its parasites in Paraguay and Bolivia. of Central and South America. CAB International,
Biocontrol News Inf 2:88 Wallingford, 635p
Molecular Identication
of Mealybugs 5
K.B. Rebijith, R. Asokan, and N.K. Krishna Kumar
Insects are the numerous life forms that have cap- has been a monumental task which calls for the
tured the attention of human beings since ancient availability of more specialists and funding. But
times. In the same context, proper classication with the dwindling interest in taxonomy and fund
and identication of life forms has been a chal- availability, the classication and identication
lenge, and a plausible method of classication of various life forms, particularly insects, has
was established by Carlous Linnaeus, a Swedish been a major challenge to the scientic commu-
botanist who published Systema Naturae in 1758. nity. With the advent of molecular biology and
However, the Linnaeus system of classication molecular tools, the identication of life forms,
was not based on evolutionary relationships including insects, has become quick, precise and
among the target groups. Later, Darwins The easy. The development of species-specic mark-
Origin of Species in 1859 changed the way life ers enables even a non-specialist to identify
forms were classied, where the identication, insects to the species level.
description and explanation of the diversity of
the organisms had come to be known as system-
atics. According to Mayr and Ashlock (1991), 5.1 Methods of Classication
systematics is the scientic study of the kinds and and Identication
diversity of organisms, and any and all relation-
ships among them; taxonomy, on the other hand, 5.1.1 Linnaean System
was the theory and practice of the classication
of organisms. It took 200 years for taxonomists Taxonomists assess the physical characteristics
to describe the 1.7 million species on the earth, that a set of species shares and selects the most
which is only 10 % of the total number of species representative species to be the type for each
estimated. In this context, identication of insects genus, and the most representative genus to be
the type of the family and so on. Individual speci-
mens are deposited in museums to serve as a ref-
K.B. Rebijith R. Asokan (*) erence for that species and genus. When new
Division of Biotechnology, Indian Institute of species are found with similar traits, they are cat-
Horticultural Research, Bangalore 560089, India
e-mail: rebijith@gmail.com; asokaniihr@gmail.com
egorized as part of a known species as a new spe-
cies, or as a new genus, depending on how closely
N.K.K. Kumar
Indian Council for Agricultural Research,
the new specimens resemble the type. The reli-
New Delhi, India ance of types results in dramatic changes if a
e-mail: ddghort@gmail.com taxonomist re-evaluates a group and decides that
some members do not belong and suggest that the ing to higher extinction rates and the introduction
group name must be changed. of invasive species of pests. Therefore, there is a
need for faster species identication and infor-
mation about their biodiversity for conserving
5.1.2 Cladistics them before they vanish from the face of the
earth. Undoubtedly, the contribution of morpho-
During the 1980s, another classication method logical taxonomy is enormous, but it also has
called cladistics, which is based on the evolution- some drawbacks, such as the following:
ary histories of organisms, was proposed. This
method is based on phylogeny, whereas the Incorrect identication due to phenotypic
Linnaean system is not. plasticity and genetic variability between dif-
ferent taxa of mealybugs
There are many morphologically cryptic taxa
5.1.3 Phylocode which are common in many groups
Morphological examination is time consum-
In this system, the genus name is removed, and ing and is often effective only for a particular
species name is shortened and hyphenated with life stage or gender (mostly in adult females in
their former genus name or given numeric case of mealybugs). As a result, many cannot
identication. be identied
Although modern interactive versions repre-
sent a major advance, the use of keys require
5.2 Shortfalls in Morphological high level of expertise that often lead to mis-
Identication identication (Hebert et al. 2003a)
Taxonomists have always looked for discon-
Mealybugs (Hemiptera: Pseudococcidae) are the tinuous character variations that could signal
major pests in a wide range of agricultural crops divergence between species. The debate on
as well as ornamental plants worldwide (Millar threshold values employing molecular identi-
2002; Miller et al. 2002, 2005). These sap- cation for interspecic divergence is also
sucking insects have been studied intensively for true in the case of morphology-based
decades because of the economic losses they identication.
cause to agriculture through direct physical dam- Early identication of new invasions is an
age to crop plants, as well as by vectoring many important aspect in preventing the spread.
plant pathogenic viruses, which in turn decreases Rapid and accurate identication of mealy-
yield quality (Meyer et al. 2008 and Nakaune bugs is not easily accomplished with conven-
et al. 2008). The family Pseudococcidae consists tional taxonomy. Taxonomy separation of
of more than 2000 described species in 270 gen- many species occurring together can be dif-
era (Downie and Gullan 2004). Current estimates cult, particularly for the nymphal stages that
suggest that the earth may have anywhere from are primarily involved
10 to more than 40 million species of organisms,
but only about 1.7 million of them have actually Hence, there is a need for an adjunct tool that
been described. It includes over 7,50,000 insects, facilitates rapid identication of species where
and it took 250 years for taxonomists to catego- molecular identication, popularly called DNA
rize all 1.7 million species, which comprise only barcoding, becomes handy. The concept of
10 % of the total species on earth (Hebert and DNA barcoding was proposed by Hebert et al.
Gregory 2005). Classifying the remaining 90 % (2003b, c) as a rapid and precise way for species
of the unidentied organisms will require more discrimination of a broad range of biological
time and expertise of taxonomists to complete specimens using a selected 658-bp fragment of
this monumental task. Economic development the 5 end of the mitochondrial cytochrome oxi-
and increased international commerce are lead- dase-I (mtCO-I) gene (Fig. 5.1).
5 Molecular Identication of Mealybugs 77
Fig. 5.1 (a) Organization of genes in mitochondrial and primer region (brown) with amino- and carboxy-
genome. (b) Arrangements of barcode region with mito- terminal spanning inside
chondrial membrane with barcode region (blue in colour)
5.2.1 Uses of DNA Barcoding cies whether abundant or rare, native or inva-
sive, engendering appreciation of biodiversity,
Works with fragments: Barcoding can identify locally and globally (Stoeckle et al. 2004).
a species from bits and pieces. When estab- Opens the way for an electronic hand-held
lished, barcoding will quickly identify unde- eld guide, the Life Barcoder: Barcoding links
sirable animal or plant material in processed biological identication to advancing frontiers
foodstuffs and detect commercial products in DNA sequencing, miniaturization in elec-
derived from regulated species (Stoeckle et al. tronics, and computerized information storage
2004). (Stoeckle et al. 2004).
Works for all stages of life: Barcoding can Demonstrates value of collections: Compiling
identify a species in its many forms, from eggs the library of barcodes begins with the multi-
and seed, through larvae and seedlings, to million specimens in museums, herbaria, zoos
adults and owers (Rebijith et al. 2012). and gardens and other biological repositories
Unmasks look-alikes: Barcoding can distin- (Stoeckle et al. 2004).
guish among species that look alike, uncover- Speeds up writing the encyclopaedia of life:
ing dangerous organisms masquerading as Compiling a library of barcodes linked to the
harmless ones, and enabling a more accurate vouchered specimens and their binomial
view of biodiversity (Asokan et al. 2011). names will enhance public access to biologi-
Reduces ambiguity: Written as a sequence of cal knowledge, helping to create an on-line
four discrete nucleotides CATG along a encyclopaedia of life on earth, with a webpage
uniform locality on genomes, a barcode of life for every species of plant and animal (Stoeckle
provides a digital identifying feature, supple- et al. 2004).
menting the more analog gradations of words,
shapes and colours (Stoeckle et al. 2004). The core idea of barcoding is based on the fact
Democratizes access: A standardized library that short pieces of DNA vary only to very a
of barcodes will empower many more people minor degree within the species, and that the
to call by name the species around them. It variation is much less between different species.
will make possible the identication of spe- Therefore, a threshold value of variation could be
78 K.B. Rebijith et al.
characterized for each taxonomic group (212 mitochondrial, is unlikely to yield data that are
%), above which groups of individuals do not balanced, universally acceptable, or sufcient in
belong to the same species, but form supra- taxonomic scope to recognize many species lin-
species taxon. Therefore, unknown individuals eages (Rubinoff 2006). Mitochondrial cyto-
could be assigned to a species level. chrome c oxidase subunit I (mtCO-I) gene
sequence is suitable for this role because its
mutation rate is often fast enough to distinguish
5.3 Targets for Molecular closely related species, and also because its
Identication sequence is conserved among conspecics and a
lack of recombination. mtCO-I sequence differ-
5.3.1 Mitochondrial DNA ences are too small to be detected between closely
related species; more than 2 % sequence diver-
Mitochondrial (mt) DNA (Fig. 5.2) has a long gence has been detected between such organ-
history of use at the species level; recent analyses isms, proving the barcode effective. However,
suggest that the use of a single gene, particularly the rate of evolution of cox1 is very slow.
Fig. 5.2 Flowchart showing steps in DNA barcoding of mealybugs, from collection to sequence deposition in iBOL
5 Molecular Identication of Mealybugs 79
Proteinase K and TNES buffer; and the samples DNA Analyzer; Applied Biosystems, USA) using
can be left for at least 1 h at 20 C during pre- PCR specic primers, both in forward and reverse
cipitation of the DNA with absolute ethanol. directions. Homology search and sequence align-
Extracted DNA can be stored at 20 C. ment can be performed employing the NCBI-
BLAST (http://blast.ncbi.nlm.nih.gov/) and
BioEdit version 7.0.9.0 (Hall 1999), respectively.
5.5.2 Polymerase Chain Reaction All the sequences generated in the respective
studies need to be deposited in the NCBI-
Polymerase chain reaction (PCR) was developed GenBank and the Barcode of Life Data systems
by Kary B. Mullis (Mullis and Faloona 1987) and (BOLD) (Table 5.2).
has radically changed molecular research and
diagnostics (Caterino et al. 2000). PCR involves
the in vitro synthesis of large amounts of DNA 5.6 Nuclear Copies
copies from a single starting molecule and of Mitochondrial Genes
employs short single strands of DNA (1830
nucleotides) called oligomers or primers (Table There is a possibility that a pseudogene is being
5.1) to select a region of specic interest from the amplied if the study encounters the following
DNA. Once the primers are annealed to the DNA, anomalies (Zhang and Hewitt 1996):
Taq DNA polymerase builds a complementary
strand extending from the primer by incorporat- More than one bands, or different bands, are
ing free deoxynucleoside triphosphate (dNTP: constantly produced during PCR
base + deoxyribose sugar + phosphate) molecules amplication.
in the reaction mix. Two primers that anneal on Background peaks or sequence ambiguities
complementary strands are used, with the Taq are constantly found when sequencing.
extending the region between them. The reaction The DNA sequence contains data which will
mixture is cycled between different temperature unexpectedly change the polymerase transla-
optima for the different stages of reaction of tion of the sequence, such as unusual frame-
denaturation, annealing and elongation. This pro- shifts, insertion/deletion or stop codons.
cess is repeated in a number of cycles (usually The DNA sequence is particularly more diver-
3040), and the DNA thus produced increases gent than expected.
exponentially (Saccaggi 2006). Phylogenetic analysis results in unusual,
unexplained or contradictory tree topology.
5.5.3 Sequence Analyses In the recent past, DNA barcoding has gained
and Submission importance in the species diagnosis of animal
species, but has some difculty with certain
The amplied products can be eluted using an insects. This is probably due to its inconsistency
extraction kit according to the manufacturers in amplifying the 5- mtCOI region; however, a
protocol, and the sequencing can be done in an total of 178 mtCOI sequences for 29 mealybug
automated sequencer (ABI prism 3730 XL species are available with the NCBI-GenBank.
Table 5.2 Maximum composite likelihood estimate of and biotypes (Shufran et al. 2000). In this regard,
the pattern of nucleotide substitution from 29 species of
sequence analyses revealed that the specimen of
mealybugs
a single species, namely Planococcus cus,
A T C G Crisicoccus matsumotoi, Phenacoccus solani
A - 10.58 2.19 2.4 and P. aceris, from various geographic regions
T 8.03 6.15 1.07
and hosts often showed substantial genetic differ-
C 8.03 29.68 1.07
ence, possibly reecting cryptic species over-
G 18.04 10.58 2.19
looked by current taxonomy classication (Park
The nucleotide frequencies are 0.367 (A), 0.484 (T/U),
et al. 2011). However, further studies are required
0.1 (C) and 0.049 (G). The transition/transversion rate
ratios are k1 = 2.247 (purines) and k2 = 2.805 (pyrimidines). in this direction to clarify these potential cases of
The overall transition/transversion bias is R = 0.443, cryptic mealybug species (Fig. 5.4).
where R = [A*G*k1 + T*C*k2]/[(A + G)*(T + C)]. Codon
positions included were 1st + 2nd + 3rd + Noncoding
5.7 Limitations of DNA
Barcoding Employing mtCOI
All the above species could be clearly differenti-
ated on the basis of the 5- mtCOI barcode (Fig. Following are the limitations of DNA barcoding
5.3), which is a valuable tool for the identica- employing mtCOI:
tion of these serious insect pests, an approach
complementing classical taxonomy. An insight Limitations in resolving species at species
into the sequence analyses revealed that the G.C boundaries in some groups where nuclear
content in the barcode region is very low (14.9 ribosomal regions are suitable.
%) and is the lowest from any insect species, mtCOI does not show much variation in
which is in total contrast to other lineages (3353 plants, except for some algae.
%) (Min and Hickey 2007; Park et al. 2011). Introgression: mtCOI is largely maternally
Interestingly, the low G.C frequency is more pre- inherited, and usually as a single copy. Hence,
dominant at the third codon (wobble position), it has one fourth the population size of other
than the rst and second position. Among various nuclear genes, has a different inheritance pat-
life forms, the G.C content (12.6 %) for tern and is more sensitive than nuclear genes
Atrococcus paludinus is the lowest value, which to population bottlenecks. mtDNA introgres-
is even lower than bacterial genomes whose G.C sion confounds the boundaries between other-
content is in the range of 1775 %. The lowest wise distinct lineages; such introgression
G.C content (1733 %) occurs in bacterial spe- between species could lead to inaccurate
cies with small genome sizes, especially the identications.
endosymbionts of insects, such as aphids Maternal inheritance: The full effect of mater-
(Andersson and Kurland 1998; Moran et al. nal inheritance on rates of molecular diver-
2008). The lower G.C content in insects such as gence in mtDNA is not predictable, and
scales and mealybugs, feeding on plant sap, a diet therefore the failure rate of DNA barcoding is
which is very decient in organic nitrogen, pos- also unpredictable. mtDNA is inherited mater-
sibly can be explained as an evolutionary adapta- nally, but not in bivalve molluscs which dis-
tion to less nitrogen and relatively less nitrogen is play double unpatented mtDNA inheritance. It
required for A.T than G.C pair. is also evident in a wide range of the taxa
DNA barcoding can be used as an acceptable infrequent paternal inheritance.
system for molecular identication of species in Low recombination: The general absence of
its distinct life stages and forms (Foottit et al. recombination will lead to the persistence of
2010; Rebijith et al. 2012), host-associated population structure long after the barriers
genetic differences (Brunner et al. 2004), dis- which created the structures are removed and
crimination of cryptic species (Smith et al. 2006) gene ow is restored. Therefore, it is not pos-
82 K.B. Rebijith et al.
99
Planococcus kraunhiae (12)
99
Planococcus lilacinus (6)
96
Planococcus ficus (4)
99 JF965419_Planococcus minor
99
Planococcus citri (9)
99
Crisicoccus pini (2)
99
Atrococcus paludinus (3)
99
Pseudococcus calceolariae (7)
99
Dysmicoccus lepelleyi (7)
99
65 Pseudococcus cryptus (5)
98
Pseudococcus com stocki (9)
99
Paracoccus marginatus (7)
99
Balanococcus takahashii (2)
99
Crisicoccus matsumotoi (19)
99
Pseudococcus jackbeardsleyi (8)
99
Pseudococcus viburni (3)
99
Dysmicoccus brevipes (3)
99
Dysmicoccus neobrevipes (15)
99
Maconellicoccus hirsutus (4)
99
Palmicultor lumpurensis (3)
99
Ferrisia virgata (2)
99
Ferrisia malvastra (4)
HM474183_Heliococcus puerariae
99
Dysmicoccus wistariae (2)
99
Pseudococcus longispinus (12)
99
Phenacoccus aceris (11)
99
Heliococcus kurilensis (7)
HM474263_Phenacoccus avenae
99
Phenacoccus solani (9)
0.01
Fig. 5.3 NJ tree with bootstrap support (1000 replicates) matsumotoi, Phenacoccus solani and P. aceris showing
showing clusters of species for mtCOI sequences. Distinct two distinct groups with >90 % bootstrap support. The
clades for 29 species of aphids can be seen in the gure, in numbers indicated in brackets represent the individuals
which four species, namely Planococcus cus, Crisicoccus analysed in the corresponding species
sible to estimate species boundaries which tent mutation rate, such as the proposed 23 %
would have been estimated from a broader divergence to correlate with species limit on a
data set. consistent basis. Speciation, uniquely driven
Mutation rate: For the DNA barcode to be by changes in mtDNA or speciation event,
used as standalone, there should be a consis- necessarily alters the mtDNA haplotypes.
5 Molecular Identication of Mealybugs 83
Fig. 5.4 Neighbour joining (NJ) showing intra-specic variation in the barcode region for four species of mealybugs,
namely Planococcus cus (a), Phenacoccus solani (b), Crisicoccus matsumotoi (c) and P. aceris (d)
Heteroplasmy: This refers to the classical barcoding initiative of the animal kingdom will
view of mitochondria functionally haploid produce about 100 million sequences and will be
with multiple identical copies. However, sin- available through GenBank.
gle nucleotide differences are common in An international database called BOLD
some species and are also abundant in some, (Barcode of Life Data system) organizes the
especially at the restriction sites. sequence data on species identication, which
Compounding genetic factors: Coinheritance was initially developed as an informatics work
factors that bias single mitochondrial inheri- bench for a single, high-volume DNA barcode
tance and most obvious are (a) mitochondrial facility. Later, the same has been selected by
selection either on the barcoding gene itself or the Canadian Barcode of Life Network (www.
on the other linked genes; (b) cytoplasmically bolnet.ca) to barcode all the eukaryotic life of
inherited bacteria like Wolbachia and some Canada, and subsequently, it has been adopted
Rickettsia which alter the inheritance factors by major barcode communities like birds,
(Rubinoff et al. 2006). shes, lepidoptera, etc. BOLD provides an inte-
Identication depends on the intra- and inter- grated bioinformatics platform that supports all
specic genetic variations. phases of analytical pathway from specimen
Difcult to resolve, recently diverged species collection to a tightly validated barcode library.
that arose through hybridization. It also provides a vehicle for collaboration
No single gene is conserved in all domains of across research communities by coupling exi-
life and exhibit enough sequence divergence ble security and data entry features with web
for species discrimination. based delivery. A copy of all sequences in
BOLD is also sent to NCBI, DDBJ, and EMBL
There are about 52 million sequence records as soon as the results are ready for public
available currently, and it is expected that the release.
84 K.B. Rebijith et al.
18S 28S
IGS ETS ITS
phosphogluco isomerase 25. phosphoenol RAPD data clearly separated the species into
pyruate carboxy kinase 26. prune 27. copper, two groups (Serce et al. 2007).
zinc superoxide dismutase 28. sodium channel 2. Seven species of mealybugs (Ps maritimus,
para locus I 29. snail 30. timeless 31. triose- Ps.vibruni. Ps.longispinus, Ps.calceolariae,
phosphate isomerase 32. vestigial 33. white Pl.cus, Pl.citri, Ferrisia gilli Gullan) were
34. wingless 35. xanthine dehydrogenase 36. identied using a Multiplex PCR based on the
yolk protein 1 and 2 37. zeste. mitochondrial cytochrome oxidase subunit
gene (Daane et al. 2011).
3. There was a slight difference in morphologi-
5.9 Limitations cal characters in the populations of
Planococcus cus, indicating that there are
May be heterozygous and present in low copy two different populations of the same species
numbers. in Tunisian vineyards. Likewise, in the molec-
Many genes contain large introns that makes it ular analyses, two separate clades were
difcult to amplify more than one exon. revealed in the neighbour-joining phyloge-
Many single-copy loci are actually are present netic tree, supporting the morphological stud-
in more than one copy. ies and suggesting there are two distinct
Pseudogenes may create problem if compari- populations of grape vine in Tunisia, which
sons are made inadvertently. might be two different biotypes (Mansour
et al. 2012).
Even with all the limitations, the molecular
identication of insects employing mtCOI is
gaining momentum, and as of now it can be an
effective adjunct tool for the integrated taxon- References
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take shape, such as the recent initiative on Andersson SG, Kurland CG (1998) Reductive evolution
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Department of Biotechnology. With the increase S, Karanth PK, Ellango R, Ramamurthy VV (2011)
in international trade on agricultural produces Molecular identication and phylogeny of Bactrocera
where the danger of introduction of invasive spe- Species (Diptera: Tephritidae). Fla Entomol
94:10261035
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a vital role in the quick identication of insect- (2004) Host associated genetic differentiation in
pests at the port of entry. As ambitiously envis- Thrips tabaci (Insecta: Thysanoptera), as determined
aged, the development and deployment of the from mtDNA sequence data. Heredity 93:364370
Caterino MS, Cho S, Sperling FAH (2000) The current
hand-held sequencer, which is supported by the state of insect molecular systematics: a thriving tower
global networked database, is going to revolu- of Babel. Ann Rev Entomol 45:154
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already described, along with the new ones. VA, Walsh DB, Zaviezo T, Almeida PP (2011)
Development of a multiplex PCR for identication of
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mealybugs (Hemiptera: Coccoidea: Pseudococcidae)
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citri, Pl.cus, P.ovae, Ps.longispinus, Ps.vibruni, 29:238259
Ph.aceris) was studied using randomly amplied Folmer O, Black M, Hoe W, Lutz R, Vrijenhoek R (1994)
polymorphic DNA-polymerase chain reaction DNA primers for amplication of mitochondrial cyto-
chrome c oxidase I from diverse metazoan inverte-
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Biology
6
M. Mani and C. Shivaraju
A few generalizations on the biology of mealy- citri (Risso). To attract adult males, the females
bugs are derived primarily from the detailed stud- emit a sex pheromone. Female mealybugs mate
ies of the work previously done on common multiple times and the number of times they mate
mealybug species. There are slight variations in also affects the egg production. Parthenogenetic
the biology of mealybugs among the species. reproduction is also observed in many mealybug
species, while in some others, reproduction is by
both sexual and parthenogenetic means.
6.1 Reproduction Most mealybug species reproduce sexually, as
well as lay eggs. However, some species such as
The mature or gravid adult female begins to grow in Phenacoccus solani Ferris, Phenacoccus parvus
size as the ovaries develop, ending at about 45 mm Morrison and Ferrisia malvastra (McDaniel)
in length and far less dorso-ventrally attened. The reproduce parthenogenically and others, for exam-
adult male is about 1.5 mm in length, with long ple, Pseudococcus longispinus (Targioni Tozzetti)
wings, a brown-coloured body and two multi-seg- and Antonina graminis (Maskell) are ovovivipa-
mented antennae that are about half the length of rous. Two different genetic systems may be found
the body. Mealybugs have the lecanoid type of the in mealybugs; the more common corresponds to a
paternal genome elimination system, where both particular type of haplodiploidy known as paternal
sexes develop from fertilized eggs (i.e., diploidy), genome elimination in which both males and
but during the early stage of development of the females develop from fertilized eggs. The male
male, the paternal half is deactivated through het- develops from a zygote containing one haploid
erochromatinization. This system suggests that the genome from his mother and one haploid genome
females would produce a male-biased sex ratio from his father, but only the maternal genome is
when alone and a more female-biased sex ratio transmitted to the offspring via the sperm, because
when grouped with other females. Mealybug repro- the set of chromosomes of paternal origin becomes
duction can be quite variable. For some mealybugs, heterochromatic and genetically inactive. Male
mating is probably necessary, although facultative mealybugs are thus functionally haploid, owing to
parthenogenesis has been reported for Planococcus heterochromatization (parahaploidy). The other
genetic system is thelytokous parthenogenesis, in
which there are no males and therefore no mating
M. Mani (*) C. Shivaraju
occurs. There are no sex chromosomes in mealy-
Indian Institute of Horticultural Research,
Bangalore 560089, India bugs; sex is probably determined by a functional
e-mail: mmani1949@yahoo.co.in haploidy/diploidy mechanism, which seems to be
dependent on the behaviour of a set of chromo- The male is holometabolic and develops
somes and not on a single chromosome. If hetero- through four stages, egg, larva (two nymphal
chromatization of an entire set of chromosomes instars), pupa and adult, while the female is
takes place during the cleavage stage of embryo- hemimetabolic and develops through three
genesis, the embryo will develop into a male, or stages, egg, larva (three nymphal instars) and
otherwise into a female. Spermatogenesis is char- adult.
acterized by inverse meiosis and the absence of
chromosome pairing and genetic recombination
where the genome of the mother determines the 6.1.1 Oviposition
heterochromatization of the inherited paternal
chromosomes in mealybug embryos. According Mealybugs may be oviparous, viviparous or ovo-
to this model, heterochromatization is controlled viparous. Although variable, the pre-oviposition
by a maternal factor, with the maternally derived period is 45 days in general. In most of the
chromosomes imprinted so that they do not suffer mealybugs like Pseudococcus longispinus,
the fate of the male chromosome. Sex determina- Ferrisia gilli Gullan, Dysmicoccus brevipes
tion in mealybugs, and consequently the sex ratio, (Cockerell) and Heliococcus bohemicus Sulc.,
is known to be inuenced by temperature and the eggs are laid by the adult female among the la-
age of the mother. The effect of the temperature or mentous secretion of the ovisac formed by the
the age of the mother on the sex ratio of the off- pores on the adult's body. Wax secreted by the
spring is attributed to a change in the ratio between numerous pores and ducts around the ovi-
the numbers of oocytes with and without the positional opening plays an important role in
maternal factor. forming a waxy sac around the laid eggs. The
6 Biology 89
ovisacs are somewhat variable in their structure, depending on the species, environmental condi-
depending upon the species of the mealybug. tions and food supply. It has been reported rang-
Several types of ovisacs have been recorded, ing from about 50 to over 800. In the absence of
varying from one covering the entire female adult males, the virgin females of Planococcus
body, to one covering only half or perhaps the minor (Maskell) do not produce eggs.
last two abdominal segments, to one which is
entirely ventral. These ovisacs are normally
deposited on the shoots, fruits, owers, leaves, 6.1.2 Incubation
underneath the bark and the cracks and crevices
of the plant stem and arms. The eggs hatch between 4 and 10 days, depend-
The eggs are oval and can be seen with the ing on the temperature and humidity. Hatching
naked eye, while the colour of the eggs varies percentage ranges from 80 to 90 %.
with the species. The normal oviposition period
of female mealybugs is 68 days, but it differs
depending upon the species and climatic condi- 6.1.3 Nymph
tions. They are in close proximity in the ovisac.
Female mealybugs are capable of mating multi- Mealybugs generally have three larval instars for
ple times on the same day and the female repro- females and four for males. Each of these stages
ductive output is unaffected by multiple resembles the previous one, except for an increase
copulations. As many as 600 eggs can be laid by in size and the amount of wax secreted. Usually,
one female, but the number of eggs varies accord- immature males are slightly longer and more
ing to the mealybug and the host species. Freshly slender than females.
laid eggs vary in colour in different species, for
example, the eggs are orange in case of
Maconellicoccus hirsutus (Green), violet in case 6.1.4 First-Instar Nymph (Both
of Nipaecoccus viridis (Newstead), yellow in Sexes)
case of most of the mealybugs like Planococcus
citri (Risso). However, the colour changes The eggs hatch into rst-instar nymphs or crawl-
slightly before hatching, and the size also varies ers and are vulnerable to insecticidal applica-
in different species. In the case of M. hirsutus, the tions. This stage usually has proportionally very
eggs are 0.340.38 mm in length and the width large legs and antennae compared with other
ranges from 0.17 to 0.20 mm. Most of the mealy- instars and often there are fewer antennal seg-
bugs are oviparous and they lay eggs in the ovi- ments than in the adult. This crawler is often
sac. But some mealybugs are ovoviviparous quite mobile and usually migrates to other hosts,
(depositing live rst instars); for example, or at least to diverse areas of the parent host.
Dysmicoccus brevipes (Cockerell) bears live There is no reliable character to distinguish
young, producing as many as 908 crawlers. These between the male and female at this stage. The
ovoviviparous females produce little or no ovisac mean duration of the rst instar is about 67
and apparently shield their young for a short time days; there is not much variation between male
by covering them with their abdomen. This char- and female instars, but it can vary under different
acter is particularly true of Puto, a genus which, conditions of temperature and humidity. When
for the most part, is ovoviviparous. Planococcus viewed from above, it seems elongate oval in
lilacinus (Cockerell) also lays the crawlers shape, but is extremely at from the side. The
directly, while Ferrisia virgata (Cockerell) lays rst-instar nymph is often free from the waxy
eggs which hatch immediately after laying. The coating which usually develops at the later stages
number of offspring produced per female varies of growth.
90 M. Mani and C. Shivaraju
6.1.5 Second-Instar Nymph (Both instar male is about 1 day, but it again varies
Sexes) according to temperature and humidity condi-
tions for different species.
The second-instar nymph emerges through a slit
in the medio-dorsum of the head and thorax of
the rst-instar skin. The legs and antennae are 6.1.7 Fourth-Instar Male
still proportionally larger compared to the adult
body and often there are fewer antennal segments The fourth instar of the male is produced in a
than in the adult. During this stage, it is possible cocoon and is known as the pupa. The third skin
to differentiate between the sexes, for by the end that is shed is again shoved to the back of the
of this instar, the males produce a lamentous sac cocoon. This instar differs from the previously
or cocoon over their bodies. The rostrum of the mentioned one; it has two longer, backwardly
male is lost after the rst moult, making the rec- directed wing pads, more antennal segments (ten-
ognition of the second-instar male quite simple. jointed, normally the same as the adult) with a
The second nymphal stage of the female is char- few pointed setae and three pairs of eyes. At one
acterized by the presence of six jointed antennae, point between this instar and the next, a break in
four pairs of cerarii in the abdominal segments the posterior part of the cocoon is formed and the
VIIX and the oral rim ducts are restricted to the fourth shed skin is pushed outside. The duration
dorsum. In the second stage, the male only has of the fourth instar/pupa is about 6 days, but it
one pair of, rarely two pairs of, cerarii and tubular again varies under different conditions of tem-
ducts of oral collar type both on the dorsum and perature and humidity.
the venter. Mean duration of the second instar is
about 67 days, but not much variation is seen
between male and female instars, though it can 6.1.8 Adult Female
be variable under different conditions of temper-
ature and humidity. The adult female is almost similar to the third-
instar female nymph. The third-instar female has
seven-jointed antennae and ve pairs of cerarii in
6.1.6 Third-Instar Nymph (Both the abdominal segments VIX, but the adult
Sexes) female has nine-jointed antennae, six pairs of
cerarii in the abdominal segments IVIX., valvu-
The second-instar skin is shed in the same man- lar opening and multilocular disc pores in the
ner as the rst, producing the third-instar nymph. venter. The fourth-instar female emerges in the
In female specimens, these instars begin to take usual manner to become an adult. The female
the normal shape of the adult. Legs and antennae nymph can be distinguished from all of the
are approximately in proportion to the body size, nymphal instars by the presence of a vulva and it
when compared with the adult and the antennal is often very large and distorted from the eggs or
segments are normally, but not always, the same nymphs which she carries in her abdomen. The
in number as in the mature forms. The duration of female nymph has various types of pores on its
the third instar in females is usually about 8 days, body; the circular multilocular and pentagonal
but it can vary under different conditions of tem- quinquelocular pores are believed to function in
perature and humidity. In the male, the second the production of the lamentous ovisac, whereas
skin is shoved to the posterior portion of the the normally more abundant trilocular pores
cocoon; the third-instar male is called the pre- function in the production of the white powdery
pupa and is usually much smaller and more elon- secretion, characteristic of mealybugs. The exact
gated than the third instar female. It also differs function of any of these pores has yet to be con-
because it has wing pads, no rostellum and more rmed and the precise morphological structure of
antennal segments. The duration of the third- the various types of tubular ducts is still unknown.
6 Biology 91
In certain mealybug species, such as Ferrisia vir- these of Pula are very dark brown or almost black
gata, it is believed that the enlarged tubular ducts in colour. Leg size is also quite variable:
produce thin crystalline rods, but the importance Antoninoides have very small, reduced legs;
of the smaller tubular duct remains a mystery. Discococcus have slightly larger legs;
The setae of the cerarii are apparently impor- Phenacoccus, Spilococcus, Chorizococcus and
tant in supporting the laments; again, in most others bear proportionately normal sized legs;
adult female mealybugs, two pairs of dorsal cavi- and the numerous Puto species bear very enlarged
ties or ostioles, as they are called, can be located and robust legs. The adult females of Antonina
submarginally. The posterior pair is on the seventh are totally without legs (apodous); on each sur-
abdominal segment and the anterior pair is appar- face of the posterior part of the trochanter, there
ently on the head. As previously mentioned, the are normally two, sometimes three, small clear
ostioles, when irritated, seem to function in the spots which are sensory in function. On the tarsal
secretion of globules of body uid, perhaps as a segments and on the claw, mealybugs normally
defensive mechanism. It is also believed that these have a pair of spatulate setae, but these are some-
dorsal openings function in the exit of honeydew times absent. The use of these setae is not known,
for consumption by ants. The laments which may but it seems logical to assume that they, in some
be seen so often on the lateral margins of many way, function in clinging to the substrate. The
mealybugs are apparently produced by the clusters vulva, which is always present in the adult
of trilocular pores surrounding the cerarii. female, is the genital opening and functions in
Other structures of the adult female are the copulation and egg laying. In some instances,
antennae, the eyes, mouthparts, legs, spiracles, more than one male is able to mate with a single
vulva, circulus, anal ring, anal lobes, body setae female at one time.
and cerarii. The antennae of mealybugs are li- The anal rings are often of various shapes in
form, with the terminal segments characteristi- different mealybugs, but they all function in waste
cally longer and wider than the preceding ones. elimination. As previously mentioned, these
There is a slight noticeable tapering of the anten- wastes, which are in many instances detrimental
nae from the rst segment to the last and the to the mealybug, are propelled long distances
antennal segmentation varies in number from two from the body by the rapid contraction of the
in some species of Antonina to nine in certain walls of the anal cavity. It is also possible that the
species of Phenacoccus. Antennal shape is anal ring serves as an exit for honeydew intended
important in the eld identication of some sub- for ant consumption and that the long setae of the
terranean mealybugs. The unusually short genic- ring serve as standards to hold the honeydew
ulate antennae of Rhizoecus, Pygmaeococcus and secretion before it is consumed by the ant.
Geococcus are easily recognized, particularly The adult female measures 2.652.80 mm in
when compared with the normal, straight anten- length and 1.751.85 mm in width, and the
nae of most other genera, which makes their eld females are wingless and as they mature, become
identication quite simple. There are often some more sessile. The female mealybug has a soft,
enlarged setae on the apical, two antennal seg- oval and attened segmented body, but the divi-
ments, which apparently act as special sensory sion between the thorax and the abdomen is not
setae. These setae differ from the other antennal distinct.
setae, being larger in diameter and more rounded
at the tip. The eyes of mealybugs are compound
and they function only in distinguishing between 6.1.9 Adult Male
light and dark.
The legs of mealybugs have a characteristic The fth-instar male refers to the adult which is
colour. For example, the legs of Rhizoecus and very complex morphologically and only the easily
Misericoccus are white, those of many species of recognized characteristics are discussed here.
Phenacoccus and Spilococcus are red, while There are two or three known primary types of
92 M. Mani and C. Shivaraju
dermal pores occurring on male pseudococcids. antennae, such as Palmicola palmarum (Ehrhorn).
The rst type is normally clustered in a circular The male has six eyes a ventral pair, a lateral
group on the posterior lateral margin of each side pair and a dorsal pair. The eyes of most males are
of the ninth abdominal segment. In Phenacoccus dark red, with the smaller lateral pair protruding
gossypii Townsend and Cockerell, there is, in more than the other two. The mouthparts of the
addition to the pair of pore clusters on the ninth male are almost nonexistent and are completely
abdominal segment, another pair on the eighth non-functional. All that remains of the mouth is a
segment. These pore groupings apparently form small circular opening found at the posterior mar-
the wax of the long caudal laments. The pores gin of the ventral part of the head. Normally,
which make up these clusters are circular with a there is one pair of wings which develope from
ve-pointed star-shaped lumen, are called stellate the mesothorax; these wings possess two longitu-
pores. Most of the stellate pore clusters also have dinal veins, the anterior one being the radius and
long slender setae, which apparently act as central the posterior one being the media. There is also a
bases for the wax of the caudal laments. It is felt complex of minute basal veins called the costal
that the long anal lobe setae of the females might complex of wing veins. There are three forms of
also function in this manner. The male caudal males, the rst form has fully developed wings
laments of Puto arcloslaphyli Ferris and Puto (macropterous); the second form has wings, but
decorosus McKenzie, herein described as new, they are reduced and nonfunctional (brachypter-
show a seta in the middle of the long waxy la- ous); and the third form is wingless (apterous).
ments. In Phenacoccus gossypii Townsend and Examples of the rst form are Phenacoccus gos-
Cockerell and Pseudococcus longispinus (Targioni sypii Townsend and Cockerell, Planococcus cilri
Tozzetti), the above characteristic was observed in (Risso), Puto lalicribellum Mc- Kenzie
the females. Other types of pores have also been Saccharicoccus sacchari (Cockerell); of the sec-
found on the males and are scattered over the body ond form is Palmicola palmarum (Ehrhorn); and
surface. These are called dermal disc pores and of the third form are Puto ambiguus (Fullaway),
name the number of loculi they contain. These P. echinalus McKenzie and P. pacicus
may vary from three to ve in number with four McKenzie. A pair of halteres is present on the
being normal, and it is further pointed out that metathorax; they are slender projections adorned
these structures are by no means at, but rather are with one or more setae at their tips. The normal
recessed into the derm, with several small protru- number of apical setae is one, but four have been
sions from the central part of the loculi. on each haltere in Puto yuccae (Coquillett). The
Body setae are also characteristic in the male. apical setae, whether four or more in number, are
Just as the normal lanceolate setae of the female re-curved in such a manner so as to hook into a
are present, so also are the thick, nger-like circular pocket in the posterior part of the
digitiform setae described by Beardsley. The lat- forewing.
ter are normally the most common of the two There are two pairs of spiracles in the male,
types; they predominate on both the surfaces of just as in the female, but in the case of the male
the body, the legs and especially the antennae. there is often a much more strongly developed
The digitiform setae, which are presumably peritreme. The legs of most males are quite simi-
sensory in function, are very similar to those on lar to those of the females, although they are usu-
the last two segments of the female mealybugs ally proportionately much longer and more
antenna. slender. The tarsus of the male has two segments
One pair of dorsal ostioles is present on the rather than the normal single segment of the
submarginal portion of the seventh abdominal female, although in most cases the additional
segment and corresponds to the posterior pair of segment is quite small and appears as a slightly
ostioles on the female. The antennae of the adult sclerotized ring. The digitules of the tarsal claws
male normally have ten segments, but occasion- of most males differ from the spatulate type of
ally there are species which have nine-segmented the female in being slender and setiform. The
6 Biology 93
claw is normally without a denticle. The external the young ones are produced over a period of up
genitalia in male is relatively simple, consisting to 2 months. On average, there are 170 offspring
of a large penile sheath, a conspicuous vertical per female during the spring and about 150 in the
slit and the penis or aedeagus. The penile sheath summer. The rst-instar nymphs are motile, but
is normally large and sclerotized with a posterior the succeeding instars are sessile and produce the
apical projection; this projection maybe of vari- felted wax covering from which a characteristic
ous shapes, but is usually broadly oval. On the excretory tube protrudes. A generation may take
ventral surface of the sheath, there is a noticeable 46 weeks, depending upon temperature and
slit from which the aedeagus often protrudes, is location.
normally tube-shaped and curves downward. The
penile sheath is apically of various shapes, from
broad as in Pseudococcus fragilis Brain, to very 6.2.2 Brevennia rehi
sharp and pointed as in P. longispinus (Targioni-
Tozzetti). The male of Puto yuccae (Coquillett), Reproduction of B. rehi Lindinger is mainly by
as well as many others of the Puto group, is quite thelyotokous parthenogenesis, viviparous parthe-
different from the normal Pseudococcid male. nogenesis and, to some extent, by oviparous sex-
The primary differences noted were eight pairs of ual reproduction. The oviposition period has
eyes, four setae on each halter, a toothed and been reported to be 24 days and the total num-
bifurcate aedeagus and a denticle on the claw. ber of eggs laid by a female varies from 42 to
The adult male has a brown-coloured body, 144, although up to 350 eggs has also been
bears two white, anal laments and is about recorded. The eggs are hyaline to yellowish-
1.5 mm in length. The development of the egg to white or pinkish-white and are elongate oval in
an adult male or female mealybug takes about shape. They are laid in groups in between the leaf
2426 days, but this can vary for different species sheath under the mealy covering; the incubation
under different climatic factors in different host period ranges from a few minutes to 39 h, with
plants. The population of the male mealybug is the hatchability of the eggs varying from 41.7 to
generally very low, compared to the females; the 93.5 %. The newly hatched nymphs are crowded
male to female ratio varies from 1:5 to 1:8, but within the waxy threads for 610 h before they
again, this is highly variable for different disperse to various parts of the same plant. The
species. pale yellowish nymph is active and the body gets
The biology of the mealybug varies with dif- covered with the waxy material on the second
ferent temperature conditions. The number of day. The nymphal period varies from 3 to 4
generations is quite variable in the weeks, where the mature females lay eggs for
Pseudococcidae; there are eight life cycles in about the same duration. The male nymphs grad-
approximately 1 year. In most of the Puto spe- ually develop wings after the rst few moults and
cies, however, only one generation occurs annu- emerge as small, active, ying insects; they are
ally and in Puto sandini Washburn, a high-altitude rarely seen and generally mate with females
species, there is one generation every 4 years. before dying a day or two after their emergence.
Adult females are wingless, robust, pink and
oval.
6.2 Biology of Important
Mealybug Species
6.2.3 Cataenococcus ensete
6.2.1 Antonina graminis
Ensete root mealybug C. ensete Williams and
The mealybug A. graminis (Maskell) reproduces Matile-Ferrero is a serious pest in southern
unisexually and up to ve generations are pro- Ethiopia. The females are viviparous and pro-
duced each year. Eggs hatch within the body and duce 253 nymphs/females. The average duration
94 M. Mani and C. Shivaraju
of the rst-, second- and third-instar nymphs was non-sexual reproduction occurs. In summer,
16.2, 18.1 and 19.7 days, respectively. The aver- both the species produce living young over a
age lifespan of the adult female is 49.9 days. The 34-week period, with the Dysmicoccus neobre-
body length and width of the adult female mealy- vipes produces 346 offspring per individual and
bugs ranged between 2.94 mm and 2.53.5 mm, D. brevipes produces 246. The rst, second and
respectively, when measured inclusive of the wax third instars or larval stages last for 1026 days,
covering. Adult female mealybugs could not sur- 622 days and 724 days, respectively. Thus, the
vive more than 3 weeks in the soil in the absence total nymphal period varies from 26 to 55 days,
of plant materials. with the average being about 34 days. The pink
form starts reproducing parthenogenetically
about 25 days after the third moult, whereas the
6.2.4 Coccidohystrix insolita gray form produces males in about a 1:1 ratio
and mating is necessary for reproduction.
The adult female has very little dorsal wax and Unmated females may live for nearly 4 months
secretes a white, waxy ovisac up to 6 times as long awaiting fertilization. There are multiple over-
as its own body, which is more typical of some lapping generations, with the life cycle of some
Coccidae. The immature stages do not secrete a being at least twice as long as that of others,
thick layer of mealy wax, the body being shiny reared under similar conditions. Adult females
yellow-green with submedian grey spots on two are plump and have a convex body, and are pink-
abdominal segments and one thoracic segment. ish in colour. Lateral wax laments are usually
The female and male nymphs of C. insolita, less than one fourth as long as the breadth of the
reared in mass on sprouted potato tubers, at 22 body and those towards the back of the insect are
and 33 Cand 6096 % RH, completed ecdysis at half as long as the body. The fecundity of the
the age of 13.92 and 14.60 days, respectively. female was 658.58 nymphs/ovisac and the pre-
The ratio of female:male was 3.24:1. Starvation larviposition period for adult females lasts for
of impregnated females had no adverse effect on about 27 days. The larviposition (giving birth to
their oviposition. The increase in age of impreg- larvae) period lasts for an average of 25 days,
nation from 5 to 40 days in females had little they give birth to about 234 progeny, but may
effect on the preoviposition and oviposition peri- produce up to 1000 crawlers. It may then live for
ods and the incubation period of eggs. Fecundity another 5 days before dying. The duration of the
is about 261 days and the longevity varies from adult female life varies from 31 to 80 days, aver-
17.66 to 51.6 days. Thirty- to forty-day-old aging about 56 days. Male pineapple mealybugs
females showed 7788 % and 9699 % reduction do not exist in Hawaii; they are observed from
in oviposition period and fecundity, respectively. Brazil. Male pineapple mealybug males are dis-
tinguished from the gray pineapple mealybug
males by the difference in the number of anten-
6.2.5 Dysmicoccus spp. nal segments. The pineapple mealybug has eight
antennal segments and the gray pineappple
There are two separate species of Dysmicoccus mealybug has ten. In addition, the pineapple
found on pineapple plants. The pink mealybug mealybug has short clavate setae on its body and
Dysmicoccus brevipes (Cockerell) which repro- appendages instead of the digitiform setae that is
duced non-sexually and the gray mealybug found on gray pineapple mealybugs. The dura-
Dysmicoccus neobrevipes Beardsley which was tion of the adult female life varies from 31 to 80
bisexual. D. brevipes reproduces non-sexually days, averaging about 56 days, where, the pre-
through a process called parthenogenesis in larviposition, larviposition and post-larviposi-
which females birth female larvae without fertil- tion periods last for an average of 27, 25 and 5
ization by males in Hawaii. In areas such as days, respectively. They give birth to about 234
Brazil, where males are present, both sexual and progeny, but may produce up to 1000 crawlers.
6 Biology 95
Another species Dysmicoccus boninensis yellow) remain motionless for 1015 min before
(Kuawana) is oviparous. The eggs are laid in a moving to the tender parts of the plant to x them-
cottony ovisac and hatch in about 10 days. The selves for feeding. The average duration of the rst
nymphal period ranges from 18 to 26 days and instar is 6.7 days and it takes about 4.4 days from
the males are necessary for reproduction. rst moult to cocoon formation in MarchApril. In
Dysmicoccus carens Williams was viviparous the cocoon, males moult three times and their
with four nymphal instars preceding the adult development inside the cocoon takes about 9 days.
stage. The life cycle of the mealybug ranges from The total nymphal period is about 20 days in
48.2 to 63.8 days when reared on CoC 671 or Co males. Thus, an adult male emerges from the
740. The mean fecundity is the lowest (117.6 cocoon after the fourth moult and is ready for cop-
crawlers/adult female) when reared on Co 6907 ulation a little after emergence. The male adults
and the highest (230.6 crawlers/adult female) on live for 13 days. Females undergo three nymphal
C 740. The longevity of males is 34 days. The instars and pass through incomplete metamorpho-
longevity of females is the greatest (32.3 days) on sis. The duration of the rst, second and third
Co 740 and the shortest (18.0 days) on CoC 671. instar was 7, 6 and 6 days, respectively. Total dura-
The mean duration of the rst instar varies from tion of the nymphal stage in females averages
4.8 days on CoC 671 to 6.1 days on Co 6806. The between 43.2 and 92.6 days at 28.9 and
duration of the second instar is the shortest 16.6 deg C, respectively, while in males it aver-
(4.1 days) on CoC 671 and the longest (5.8 days) ages to 25.4 days at 26.5 C. The total lifespan,
on Co 7704. The duration of the third instar was from the egg stage to the end of the adult stage,
the shortest (5.1 days) on CoC 671 and the lon- averages about 76.2154.6 days in females as
gest (6.1 days) on Co 6806. The duration of the opposed to 1947 days in males. The male:female
fourth instar ranged from 13.5 days on Co 6907 sex ratio is 1:1.87. Adult females are apterous with
to 15 days on Co 7704. two long prominent waxy laments at the poste-
rior end and with a lot of waxy or glossy hair over
the body which is covered with white waxy pow-
6.2.6 Ferrisia virgata der. They have fairly long, dark stripes on the dor-
sum of the posterior end of the body.
Reproduction in F. virgata (CklI.) is by both sex-
ual and parthenogenetic means, but the latter is
more common in this species. The courtship lasts 6.2.7 Ferrisicoccus psidii
for 130 min and the copulation time ranges from
15 to 20 min. The males prefer young adult In F. psidii Mukhopadhyay and Ghose, the duration
females and mating occurs only once during the of the rst nymphal stage ranges from 4 to 11 days.
lifespan of the female. The longevity of the female The second- and third-instar female nymphs com-
is about 50 days and the fecundity ranges from 300 plete their moulting at the age of 15.5 and
to 700 eggs per female. The eggs are deposited in 21.35 days, being 66.8 and 60.0 % at the age of
groups, rarely singly and are usually concealed 1317 and 1922 days, respectively. In he second,
under the body. They are oval and buff to light yel- third and fourth instars, males moult at the age of
low in colour. Since the eggs are laid in an 13.28, 14.71 and 18.69 days and are around 69.4,
advanced stage of embryonic development, they 62.4 and 68.5 % developed at the age of 1114,
hatch soon after the oviposition in a very short 1316 and 1720 days, respectively. The colour of
time and are therefore seldom seen. Under normal the crawlers and all the nymphal instars of females
conditions, the egg hatching percentage is about are rosy, creamy pink, pinkish chocolate and choc-
95 %. The egg period lasts about 28.14 min. Pre- olate; waxy dusts are found on their dorsum, the
oviposition, oviposition and post-oviposition peri- quantity progressively increasing with the progress
ods are 6.4, 8.1 and 1.5 days, respectively. Upon of their development and the stage. Nymphal
hatching, the crawlers (0.34 mm long and light instars of females secrete 78 pairs and 13 pairs of
96 M. Mani and C. Shivaraju
marginally waxy tassels, mostly abdominal. All the ized by well-developed wing pads and lasts for
instars of females and the second instar of males about 57 days. The duration of development,
secrete a tubular and waxy anal process. from egg to adult in case of the female and the
male, is 30.3 and 28.7 days, respectively. The
pre-ovipositional period ranges from 6 to 7 days
6.2.8 Kiritshenkella sacchari and the ovipositional period ranges from 7 to 9
days, while the fecundity ranges from 426 to 573
The eggs of K. Sacchari (Green) are laid in a eggs. Adult males are orange coloured, minute
chain containing nearly 120 smooth eggs beneath and very active. The longevity of adult females
the abdomen of the female. The incubation period range between 13 and 16 days and for males
is 14 h when the average temperature is 27.8 C between 3 and 5 days.
and humidity is 63 %. Freshly emerged nymphs
remain beneath the abdomen of the mother for a
short while, after which they turn restive and 6.2.10 Nipaecoccus viridis
move about to settle in the vicinity of the mother.
The total life cycle is completed in 18.6 days dur- Adult females of N. viridis (Newstead) are rather
ing April. large and have black or purplish bodies. They
appear to be at, having short laments around
the margin, whereas the males are winged with
6.2.9 Maconellicoccus hirsutus long antennae. The eggs are laid in clusters and
enclosed in a protective, cottony mass. A female
Parthenogenesis was the main mode of reproduc- lays about 300500 eggs in its life time; the eggs
tion in M. hirsutus (Green), but sexual reproduc- are purple in colour and hatch in 1020 days and
tion was also observed. Freshly laid eggs are soon envelope themselves in the uffy material.
translucent and yellowish or light orange in The nymphs are amber coloured with whitish,
colour. They are elongated and oval in shape. As waxy coating around the margins. Female nymphs
the incubation period advances, the translucent moult thrice and complete their life cycle in 68
eggs become pinkish in colour before hatching. weeks, while the males moult four times and after
The incubation period varies from 5 to 7 days; passing through a pre-pupal stage, emerge as
and the hatching percentage of the eggs is about winged adults. The average developmental peri-
90 %. The rst-instar nymphs are usually yellow ods of males and females are 18.19 and 16.19 days,
to orange in colour with reddish compound eyes. respectively. The average pre-oviposition, ovipo-
The neonate larvae are oval in shape and are sition and fecundity are 7.33 days, 8.33 days and
highly mobile; during this stage, males and 176.33 eggs, respectively.
females are indistinguishable. The duration of the
rst-instar nymph lasts for 79 days. The body is
pinkish in colour with white, thin and waxy 6.2.11 Paracoccus marginatus
secretions on the body. The duration of the
second-instar nymph lasts for 68 days; at the A single female of Pa. marginatus Williams and
end of second instar, the female nymphs moult as Granara de Willink is known to lay about 230
usual, like the previous instars, but the males 400 eggs in an ovisac. The ovisac, developed
secrete a cottony puparia around their body. The ventrally, is three to four times the length of the
duration of the last instar of the female nymph body and is entirely covered with white wax. Egg
lasts for 810 days. The third-instar male nymphs laying usually occurs over a period of 12 weeks.
are recognized by denuding the puparia, distin- The eggs are greenish-yellow and egg hatching
guished by the presence of two small wing buds. occurs in about 10 days. The males have four
This instar lasts for 12 days, with an average of instars; the rst-instar nymphs are called crawlers
1.4 days. The last instar male nymph is character- and the duration of the rst-, second-, third- and
6 Biology 97
fourth- instar in the male nymph at 25 C was 6.5, 9.08 days, 67.42 eggs per female and 4.57 days,
6.6, 2.4 and 41 days, respectively. The fourth respectively. The longevity of the adult females
instar is produced in a cocoon and is referred to range from 47 to 55 days and the species repro-
as the pupa. Adult males tend to be pink in colour, duces parthenogenetically.
especially during the pre-pupal and pupal stages,
but appear yellow in the rst and second instar.
The duration of the rst-, second- and third- 6.2.14 Phenacoccus herreni
instar of the female nymph in the mealybug was
6.5, 5.5 and 5.2 days, respectively at 25 C. The Phenacoccus herreni Cox and Williams is a sex-
species is known to reproduce both sexually and ually reproducing mealybug. The rst instar
parthenogenetically. Males have longer develop- nymphs (crawlers) complete their development
ment time (2730 days) than females (2426 in an average of 7.7 days, the second instar aver-
days). The mean longevity of adult males and ages to 5.1 days and the third averages to 5.6 days.
females was 2.3 and 21.2 days respectively. The Adult females live for an average of 24.8 days,
adult female body is greenish-yellow, dusted with the oviposition period averaging about
with mealywax and not thick enough to hide the 18.4 days. The males passed through two
body colour without discrete bare areas on the nymphal instars, a pre-pupal stage and a pupal
dorsum and with many short waxy laments stage before becoming adults; these average at
around the margin of the body. around 7.5, 6, 2.8 and 3.1 days, respectively. The
ratio of females:males was 3:1. Reproduction
was exclusively sexual; oviposition begins 3 days
6.2.12 Phenacoccus aceris after pairing and the females deposit an average
of 773.6 eggs each. Initially, the crawlers are
Female Ph. aceris Signoret starts laying 200500 found mainly on the growing point of the plant,
eggs in a cottony ovisac during a 2-week period. from which they disperse down the stalk, settling
The eggs are oval and lemon-yellow in colour. nally on the lower surface of the leaves.
The ovisacs contain up to several hundred eggs.
The rst instar nymph is lemon-yellow, but with
bright red eyes. The nymphs gradually disperse 6.2.15 Phenacoccus madeirensis
to nearby plant tissues. Soon after they begin
feeding, they develop a granular white waxy cov- The total duration of development of the female
ering with laments at the caudal end, which is Phenacoccus madeirensis Green is 30 days at
typical of mealybugs. The adult female has a sage 25 C, 46 days at 20 C and 66 days at 15 C. The
green body, which is visible through the white, developmental time of males was 39 days lon-
waxy coating. The tails on the caudal end of the ger than females. Adult longevity at 25 C for
mealybug are shorter and the colour of the body males and ovipositing females was 3 and 20 days,
ranges from greenish to pale purple. respectively. Females at 20 C produced the high-
est number of eggs (500 eggs/female).
tion of the egg stage, rst instar (crawler), sec- produces a sac with a cottony covering protrud-
ondfourth instars and the adult stage averages ing from the anal end of the body. The incuba-
about 8.0, 4.5, 4.1, 4.2, 5.2 and 20.2 days, respec- tion period of the eggs is 6.6 days; the female
tively. The mean generation time is 28.48 days. nymphs moult three times, while the males
moult four times. The freshly emerged rst
instar nymphs are oblong in shape. The average
6.2.17 Phenacoccus peruvianus duration of the rst-, second- and third-instar
nymphs is 4.8, 5.6 and 6.4 days, respectively,
Adult females are elongate oval in shape, are with the total nymphal duration being 16.8 days
greyish with a green tinge and covered in a thin in females. The rst- and second-instar nymphs
layer of white mealy wax. They lack marginal are pale yellow in colour and oblong-shaped.
and caudal wax laments, which are well devel- During the third instar, a white waxy substance
oped in other mealybug species. No males are covers the dorsal body surface. The adult
observed in the case of Ph. peruvianus Granara female is oblong in shape, light to dark yellow
de Willink and they reproduce parthenogeneti- in appearance and is wingless. The pre-
cally. Eggs are laid in the highly conspicuous oviposition, oviposition and post-oviposition
white, waxy and elongate ovisacs that form dense periods are recorded as 4.3, 8.0 and 2.7 days,
groups on the undersides of the foliage and on the respectively. The female adult survives for
stems. The nymphal instars are pale orange in 15.5 days and the entire lifespan lasts about 31
colour. days. A pair of dark spots on the thorax and
three pairs on the abdomen forming two longi-
tudinal stripes are noticed. Male mealybugs
6.2.18 Phenacoccus saccharifolii have two nymphal instars. The mean duration
of the rst and second instar is 4.0 and 5.3 days,
The female Ph. saccharifolii Williams secretes respectively. At the end of the second nymphal
the ovisac probably from the accessory glands instar, the males construct the puparia. The
one or two days prior to oviposition. The gravid pupal duration ranges from 6 to 7 days. The
female lays about 700 eggs in a single ovisac in total development of the female is complete in
batches. By the time the last batch of eggs is laid, about 26 days, while that of male takes about
the body of the female is raised to a vertical posi- 18.33 days. The sex ratio is 1:1.29.
tion with the anterior end attached to the substra-
tum by the oral bristles. The female dies soon
after oviposition. The incubation period lasts for 6.2.20 Planococcoides njalensis
56 days. The total life cycle is completed in
2528 days. This mealybug, Pl. njalensis (Laing) is biolog-
ically variable. Some of the forms exhibit a
strong parthenogenetic habit, so that males are
6.2.19 Phenacoccus solenopsis not necessary, whereas the others require males
for reproduction. Fecundity is very low, aver-
Phenacoccus solenopsis (Tinsley) is observed aging only about 36 young ones per female
to be ovoviviparous; the adult female is capable over a 20-day adult lifespan. The eggs hatch
of reproducing only if she mates with a male. P. within a few moments of being laid. There is
solenopsis lays about 500 eggs in the ovisac; no ovisac, only a few thin laments being pro-
they are minute, oval in shape and light yellow vided by the female for temporary protection
in colour. The eggs are smooth, translucent of the young. The life cycle is completed in
oblong in shape with tapering ends. It retains its about 42 days and there are about eight genera-
eggs in the body until they are ready to hatch. It tions in a year.
6 Biology 99
The duration of the rst and second instar(female), called a pre-pupa, is contained in a cocoon that
the second instar (male), the third instar(female), begins forming toward the end of the second
the third instar (m) and the fourth instar(M) are instar. The fourth stage of the immature male is
6.6, 7.2, 6.4, 6.9, 2.6, 5.9 days, respectively. The the pupa; it is elongated and light reddish-brown.
females take 30.8 and males take 27.5 days, At 30 C, a generation may be produced in 2729
respectively, to complete the development cycle. days.
Fecundity is about 180 eggs and the male:female Adult females and males emerge at the same
ratio ranges from 1:1.54 to 1:2.75. time, from late June to mid-July for the rst
(overwintering) generation and late August to
mid-September for the second (summer) genera-
6.2.26 Pseudococcus comstocki tion. Adult females are present for a total of 46
weeks and oviposit for about 1 week after mat-
Pseudococcus comstocki (Kuwana) is known as a ing. The males survive for only a few days after
pest of pome and stone fruits and certain orna- emerging. The overwintered eggs hatch from
mental plants. There are two, three, or even four mid-April through May and the nymphs (crawl-
generations per year, depending to some extent ers) migrate from the oviposition sites to their
on climate. Generally, there is little overlapping feeding sites on terminal growth and to the under-
of broods until late in the season. Overwintering sides of the leaves of trees and shrubs. This hatch
is in the egg stage within the cottony ovisac. is completed by the petal-fall stage of pears. The
Overwintering eggs are laid as early as October, nymphs that hatch from these overwintered eggs
even in the milder climates, such as that of cen- are active from roughly early May to early July.
tral California. The overwintering eggs generally As the nymphs approach the adult stage, they
start hatching in late spring; they are elliptical tend to congregate on older branches at a pruning
and bright orange-yellow in colour. They are laid scar, a node, or at a branch base, as well as inside
in jumbled masses along with the waxy lamen- the calyx of pears. The second (summer) genera-
tous secretions in protected places such as under- tion nymphs are present from about mid-July to
bark crevices, near pruning cuts and occasionally mid-September.
in the calyx of fruits. The summer-generation
eggs are laid from mid-June through late July and
the overwintering ones from mid-August to 6.2.27 Pseudococcus cryptus
October. The summer-generation eggs have an
incubation period of about 11 days; the females Egg development time in Ps. cryptus (Ps.citricu-
produce an average of 200300 eggs, although lus) Hempel decreases with increasing tempera-
some individuals produce up to 700. The young ture and ranges from 2.4 days at 16 C to 1.0 days
females develop through three instars, after at 28 C. The total development time of nymphs
which they are capable of being fertilized and decreases from 54.9 days at 16 C to 17.4 days at
oviposition follows after 1015 days. The rst- 28 C and 19.3 days at 32 C. P. cryptus shows an
and second-larval instars of the female and the ovoviviparous reproductive behaviour and hence
male mealybug are virtually indistinguishable. the egg period is combined with the rst-instar
They appear similar to adult females, except that nymph. By tting linear models to the data, the
they are smaller, more oval-shaped, lack the long lower developmental threshold temperatures for
body laments and are more orange-yellowish the egg to the rst nymphs, second nymphs, third
because they have a lesser amount of wax cover- nymphs and from the egg to the third nymphs are
ing them. The rst-instar female crawler is at calculated as 8.7, 12.8, 13.1 and 12.1 C, respec-
and pale yellow but become darker over time. tively. The thermal constants are 198.6, 84.7,
The second and third instar females are similar in 69.8 and 296.3 degree-days, respectively, for
appearance, but become progressively browner each of the above stages. The non-linear model
and redder. The third instar of the immature male, based on a Gaussian equation, used to predict the
6 Biology 101
relationship between development rate and tem- similar to that of the other mealybugs. At the end
perature, is well described for all the stages. In of the second instar, the males produce cocoons
addition, the adult longevity decreases from (pupa) over their bodies. The third moulting takes
80.4 days at 16 C to 31.3 days at place within the cocoon; the fourth instar, also
32.0 C. Furthermore, the pre-oviposition and known as pupa, is characterized by well-devel-
oviposition periods show a pattern similar to that oped wing pads. Only males pupate and develop
of longevity. Overall, P. cryptus has a maximum into adult males. The male development, includ-
fecundity of 111 eggs per female at 28 C, which ing the nymphal and pupal stages, ranges from 18
declines to 102.7 eggs per female at 32 C. to 20 days with a mean of 19.10 days. The adult
female mealybugs are very sluggish and are simi-
lar to the nymphs. The male mealybugs are rare,
6.2.28 Pseudococcus jackbeardsleyi tiny and active. They have a pair of wings and two
long waxy caudal laments at the posterior end of
The Jack Beardley mealybug Ps. jackbeardsleyi the abdomen, similar to the other mealybugs.
Gimpel and Miller is oviparous and lays yellow- They are y-like insects, do not feed and die soon
ish eggs. The eggs are laid in a mass within a after they mate. The females complete the life
loose, thin and waxy sac behind their abdomen. cycle in 2529 days, with a mean of 26.20 days
The ovisac is a little elongated and the number of and while the males complete their development
eggs laid by a single mealybug ranges from 650 to in 2326 days, with a mean of 24.40 days. There
900 with a mean of 775.60. They are in close is a variation in the developmental period from
proximity within the white ovisac. Freshly laid eggs to adults in the mealybugs, depending on the
eggs were yellowish, smooth and oval with weather and host plants.
slightly tapering ends, but they turn a darkish yel-
low before they hatch. The incubation period is
58 days, with a mean of 5.37 days at 25 1.8 C 6.2.29 Pseudococcus longispinus
and 7085 % RH. The nymphs remain in the egg
sac for a day or two after hatching, before crawl- The female Ps. longispinus (Targioni-Tozzetti)
ing about the plant in search of food. Newly produces around 200 live young (which she
hatched mealybugs (crawlers) are quite active. deposits under her body) over a 23-week period.
The crawlers, once they begin feeding, secrete a During summer, the life cycle is completed in
white, waxy material that covers their body and around 6 weeks and in about 12 weeks in winter.
produces approximately 34 leg-like laments Long-tailed mealybugs produce live young, but
around the perimeter of the body. The nymphs are do not produce an ovisac. The eggs are straw yel-
light yellow and six-legged with oval, attened low at rst, but deepen in colour before hatching.
and smooth bodies. The females change only The eggs (20240) may hatch as soon as they are
slightly in appearance, except for growing in size laid, giving the impression that the young are
to about one sixth or one fourth of an inch when born, rather than hatched. The crawlers are at,
fully grown. The females of this species have oval, light yellow in colour and six-legged insects
three larval stages (or instars); similar to the other with smooth bodies. Soon after beginning to
mealybugs, the male and female nymphs are feed, they exude a white, waxy covering over
indistinguishable in the rst instar, but by the end their bodies. The differentiation between the
of the second instar, it is possible to differentiate male and the female begins only after moulting.
between the sexes. Female mealybug nymphs are The male nymphs stop feeding near the end of the
similar to that of adult female mealybugs, except second stage and migrate towards a protected
the latter are larger in size. The females become place where they secrete waxy cocoons in which
adults after the last moult; the female nymphal they complete their development. The females go
period ranges from 18 to 21 days with a mean of through three stages to adulthood, but change
20.82 days. The males have four nymphal instars, little in appearance.
102 M. Mani and C. Shivaraju
6.2.30 Pseudococcus mandio has ve. The pre-oviposition period is 12 days and
the eggs laid by the gravid females are observed on
At 25 C, 80 % RH and constant light, the female the underside of their abdomens. Fecundity ranges
Ps. mandio Williams has three nymphal instars, from 200 to 300 eggs per female; they are creamy
with average durations of 9.2, 5.7 and 6.5 days, yellow and covered with mealy powder. The rst-
respectively and their average lifespan is 17.8 days. instar nymph is cream-coloured and after the rst
The males have two instars, which last an average moult, the cream-yellow colour changes to light
of 8.9 and 5.2 days, with the average pre-pupal and pink. The nymphs feed together for some time and
pupal periods of 12.5 days, and an average adult a few days before the second moult, the nymphs
lifespan of 2.1 days. The pre-oviposition period developing as males spin a cocoon. Such male
lasts 4.7 days and each female lays an average of nymphs descend from the stalk of young plants and
302.2 eggs. The average incubation period is pupate in the leaf sheaths, while the female nymphs
3.8 days, with 99.4 % eclosion. The life cycle from remain feeding on the leaves. This stage lasts for
oviposition to adult emergence is 25.2 days for 67 days. The pre-pupa appears pink in colour and
females and 30.4 days for males. lasts for 2 days. In the case of males, the pupa is
distinguished from the pre-pupa by the presence of
wing pads. The pupa moults once again to attain the
6.2.31 Pseudococcus maritimus adult form; the adult male is a small, delicate and
alate insect with a reddish-pink body. The lifespan
Both sexes of Ps. maritimus (Ehrhorn) are capable of the male is only days. In the case of females, the
of mating multiple times on the same day and on light pink second-instar nymph is covered by mealy
sequential days. Median times between copula- secretion and this stage lasts for 6 days. Third instar
tions are short (<10 min) on the rst day that the nymph is light pink and covered with copious secre-
males are presented with the females, but tend to tions of wax. The yellowish-brown adult female is
increase with sequential copulation events. densely covered with wax, very sluggish and sel-
Unmated females live for up to 19 weeks, as mat- dom moves away from the spot of feeding. The
ing and oviposition result in reduced longevity. lifespan of the adult female, including the pre and
The eggs that are laid are yellow to orange in post-gestational period, is 2527 days.
colour and are within an egg sac. The crawlers are
yellow to orange-brown in colour. There is a stron-
ger winter dormancy in the egg and crawler stages, 6.2.33 Pseudococcus viburni
so that the seasonal development is attuned to a
deciduous host. Overwintering usually takes Pseudococcus viburni (Signoret) (=Pseudococcus
places ordinarily in crawlers and unhatched eggs obscurus Essig; Ps. afnis (Maskell)) has four or
in the loose cottony ovisac. With the rst warm ve generations per year on citrus, depending on
weather of early spring, these nymphs move to the the climate. It overwinters in all stages, with mod-
swelling buds and feed on the tender shoots; on erate retardation from cold weather. Each female
reaching maturity, they begin to oviposition around deposits up to 500 eggs during the rst 12 weeks,
June or July. The mature females tend to move to which accumulate in a loose caudal egg sac. They
the trunks or protected crevices of the rough bark hatch in about 8 days under summer conditions
to oviposition. It is this brood which, by feeding on and maturity is attained in about 42 days, fol-
the leaves and the fruits, causes the damage. lowed by oviposition after several weeks.
Parthenogenesis and sexual reproduction are the The female R. iceryoides Green lays eggs only
common modes of reproduction in this species. after fertilization. The pre-oviposition period
While the female undergoes four instars, the male lasts for 79 days and the oviposition lasts for
6 Biology 103
7 days. About 500 eggs are laid in the white may be found swarming from below the mother,
ovisac and the fecundity averages to about 800 which tend to give an impression that the mealy-
eggs. The incubation period is 6 days. Females bug is viviparous. The crawler extricates itself
moult three times, while males moult four from one end of the egg, the eggshell sticking on
times to become adults. The females take its posterior end. The rst-instar nymphs are tiny,
2030 days and males 1825 days to complete transparent and pink in colour and very active.
the life cycle. This stage lasts for 5.3 days. During the second-
instar nymphal period, the body grows in size and
the antennal length increases to 0.36 mm due to
6.2.35 Rastrococcus invadens the addition of a segment. The stage is completed
in 4.83 days. The duration of the third-instar
The mealybug, R. invadens Williams completes nymph is 17.2 days. Ovarian development is
eight generations in a year. The female and completed in 13.8 days while one generation is
male nymphs complete development in 34.67 completed in 54.7 days.
and 38.16 days, respectively, during winter at
1521 C and in 24.6332.67 days or 27.63
36.18 days at 1833 C during February to 6.2.37 Trionymus haancheni
November, the optimum being during June
July at 2731 C in females and during May The adult female Trionymus haancheni McKenzie
June at 2733 C in males. The male:female is quite small, but is visible to the eye without
ratio ranges from 2.13:1 to 3.3:1. The maxi- magnication, reaching a length of approxi-
mum pre-reproductive period and oviposition mately one fth of an inch (5 mm). The eggs are
period and minimum fecundity are 2029 days, laid in loose, cottony wax. These cottony egg
3445 days and 165 (145175) in nymphs, sacs are usually laid on the lower part of the
respectively, in winter. The minimum pre- plant, close to the roots and were also observed
reproductive period and the maximum fecun- under the leaf sheaths of the plant. A single
dity are 1418 days and 204 (180235) days in female can lay as many as 256 eggs in a single
nymphs, respectively, in September and the ovisac during a week. Reproduction occurs asex-
minimum oviposition period (2835 days) ually in the absence of males. The eggs are pink-
occurs in April. red and not visible to the naked eye. Eggs hatch
producing the crawlers (the most mobile nymphal
stage, which disperse to nd suitable sites for
6.2.36 Saccharicoccus sacchari feeding on plant sap). The crawlers can also be
transported to other plants by wind, people, or
In the largely parthenogenetic mode of reproduc- animals; they develop through several successive
tion, alate males are not uncommon, though nymphal instars that resemble small adults, each
apterous forms are also observed in the case of of which have legs and can actively move, until
Sa. sacchari (Cockerell). With a pre-oviposition the mature adult stage is reached and the cycle
period of 13.83 days, a single female is capable repeats. The number of generations in Idaho is
of depositing nearly 1,000 eggs. The eggs are still unknown but all the instars can be found at a
smooth, yellowish, cylindrical, with both ends single time on a plant host. The number of gen-
rounded. A single such batch may contain a max- erations is not known, but all the stages have been
imum of 262 eggs. The nymphs hatch within found co-existing on infested plants. Coupled
34.15 h and before hatching, the eggs become with a short generation time, the ability to repro-
soft and elongated. Sometimes, no eggs may be duce asexually can allow mealybug infestations
noticed and only orange coloured tiny crawlers to increase quickly to damaging levels.
104 M. Mani and C. Shivaraju
6.3.1 Geococcus citrinus Kuwana They are bisexual and can be ovoviviparous or
viviparous. The favourable period for their repro-
This is a bisexual species. The females lay the duction is around AugustOctober, with 30 nymphs
eggs in masses or chains. These eggs are pearly per female mealybug. The nymphs develop into
white, translucent and elongate oval in shape. both male and female adults. The males are charac-
The average incubation period is 12.20 days. In terized by one pair of wings, are shorter in size than
Gococcus citrinus, a single female is known to the females and occur in very few numbers. The
lay about 113188 eggs, with an average of females are plump, convex in shape and covered
128.2 eggs. There are three nymphal instars for with white waxy mealy substances. They develop
the females; the nymphs are elongate oval, white by undergoing three nymphal instars, while the
in colour and cluster on the roots to feed. The males undergo four growth stages. The life cycle of
duration of the rst-, second- and third-instar the females takes 33.543.7 days, while that of the
nymphs lasts for an average of 7.3, 5.6 and males take 29.339.5 days.
5.8 days, respectively. The size of the nymphs
increases with the instars. In males, the pupal
stage lasts for 5 days. The adult females are 6.3.3 Cataenococcus ensete
elongate oval, white and wingless with a seg-
mented body. The females live for about Adults of enset root mealybug C. ensete (Williams
12.78 days. The adult males are light brown in and Matile-Ferrero) are viviparous and produce
colour and have only one pair of narrow, elon- 156383 nymphs, and their total lifespan is 94113
gated and opaque wings with a round outer mar- days. This species has three nymphal stages; the
gin. The males live for a maximum of 5 days. development of the nymph to the adult mealybug
Unlike other mealybugs, G.citrinus nymphs and takes 54 days on average and the lifespan of the
adults do not produce honeydew; hence usually adult root mealybug is 50 days. The average dura-
these are not associated with ants. The total life tion of the rst-, second- and third-instar nymphs
cycle of G. citrinus and Geococcus coffeae is 16.2, 18 and 20 days, respectively. The average
Green are 28.8 days and 33.8 days, respectively. lifespan of the adult female is 50 days.
In case of Rhizoecus hibisci Kawai and
Takagi, the eggs are laid in white, loose, waxy,
elongate ovisacs which are about 2 mm long and 6.3.4 Rhizoecus amorphophalli
can easily disintegrate when disturbed. Each ovi-
sac contains up to 80 eggs, which usually hatch Rhizoecus amorphophalli is the noxious pest
within 24 h. Nymphs (immature stages) are infesting the stored tubers of elephant foot yam,
creamy-white. They closely resemble the adults, taro and tannia. The ovoid, pale white eggs are laid
but are signicantly smaller. Adult females are in clusters inside the egg sac and turn light brown
creamy-white, elongate and covered in a pow- on hatching. The average length and breadth of the
dery wax that is deposited on the roots and the eggs are 187.80 m and 102.50 m, respectively.
soil; these deposits are often the rst sign of After eclosion, the rst-instar larvae (crawlers)
infestation. Adults and nymphs feed on the roots moved out of their ovisac, actively searching for
of the host plants, but may also be found within suitable feeding sites on the tubers. The crawlers
the root ball and on the inner surface of the plant are oval and semi-translucent with three pairs of
container. The males are extremely rare and are legs and paired eyes, measuring 183 m in length
unlikely to be seen. There can be several over- and 98 m in width. They prefer to hide out in the
lapping generations throughout the year and crevices or depressions of the tubers and on settle-
their numbers can multiply rapidly under favour- ment produce mealy substance to create waxy
able conditions. laments over their body. The rst instar lasts for
6 Biology 105
47 days. The second-instar larva is 270 m long Bartlett BB (1978) Pseudococcidae. In: Clausen CP (ed)
Introduced parasites and predators of arthropod pests
and 124.74 m wide and is semi-translucent with
and weeds: a world review. USDA-ARS, Washington,
a shiny body. The second-instar larva lasts for an DC, pp 137170
average of 4.82 days and before undergoing next Chong JH, van Iersel MW, Oetting RD (2004) Effects of
moulting, they settle either on the previous site or elevated carbon dioxide levels and temperature on the
life history of the Madeira mealybug (Hemiptera:
on another suitable site on the tuber and cover
Pseudococcidae). J Entomol Sci 39:387397
themselves with mealy laments. The third instar Coumara Radja N (1985) Studies on the biology and con-
is relatively larger, measuring 429.47 m in length, trol of rice mealybug, Brevennia rehi (Lindinger)
193.2 m in width, and the duration of this instar (Pseudococcidae: Hemiptera). MSc thesis, Tamil
Nadu Agricultural University, Coimbatore
is 46 days and the sex differentiation is obvious at
Daane KM, Rodrigo PP, Almeida, Bell VA, Walker JTS,
the end of this stage. The female nymphs moult Botton M, Fallahzadesh M, Mani M, Miano JL, Sforza
normally, but the male instar produces a cottony R, Walton VM, Zaveizo T (2012) Biology and man-
puparium around its body.. The pupal stage lasts agement of mealybugs in vineyards. In: Bostanian NJ
et al (eds) Arthropod management in vineyard pests,
for an average of 2.50 days and the males trans-
approaches, and future directions, Springer Science+
form into winged adult forms. They undergo a Media B.V., Dordrecht, pp 271307. doi
radical change during their life cycle the wing- 10.1007/978-007-4032-7-12
less nymphs transform into winged adults. The Jayanthi R (1986) Mealybugs. In: David H, Easwaramoorthy
S, Jayanthi R (eds) Sugarcane entomology in India.
adult female body is oval, whitish, wingless and
Sugarcane Breeding Institute, Coimbatore, pp 259275
sparsely covered with white mealy substance. The Kamariya NM, Patel VN (2011) Biology of mealy bug,
length and width of adult females are 867.19 m Phenacoccus solenopsis (Tinsley) on cotton. J Cotton
and 368.88.78 m, respectively. After mating, the Res Dev 25(1):115118
Kim SC, Song J-H, Kim D-S (2008) Effect of temperature
adult females secrete an ovisac of white waxy sub-
on the development and fecundity of the Cryptic
stance in about 714 h and egg laying begins 37 h Mealybug, Pseudococcus cryptus, in the laboratory.
after this process. The eggs are laid in a bead- J Asia-Pacic Entomol 11(3):149153
shaped pattern, but later they are found in a disar- McKenzie HL (1967) Biology (Life history, Annual
Successions) morphology and classication.
rayed and scattered manner under mealy covering.
Mealybugs of California. University of California
Oviposition is completed in about 38 h with a Press, pp 2427, 3640
maximum fecundity of 79 eggs and the females Milena Varela A, Belloti AC (1981) Some aspects of the
are not able to survive more than 4 h after egg lay- biology of and observations on a new mealybug of
cassava Phenacoccus herreni (Homoptera:
ing. At 32.2235.10 C and 5565 % humidity, all
Pseudococcidae) in Colombia. [Spanish]. Revista
the eggs hatch after about 69 days of incubation. Colombiana de Entomologia 7(1/2):2126
Mishra D, Mukhopadhyay AK, Pramanik A (2004)
Biology of the mealybug Phenacoccus bengalensis
Pramanik and Ghose, 1999 (Homoptera:
References Pseudococcidae). Uttar Pradesh J Zool
24(2):155159
Addis T, Azerefegne F, Blomme G, Kanaujia K (2008) Mukhopadhyay AK (2005) Study on the biology of the
Biology of the enset root mealybug, Cataenococcus mealybug, Ferrisicoccus psidii. Ann Plant Protect Sci
ensete and its geographical distribution in Southern 13(1):239240
Ethiopia. J Appl Biosci 8(1):251260 Nakahira K, Arakawa R (2006) Development and repro-
Alam S, Karim ANMR (1981) Rice mealybug, Brevennia duction of an exotic pest mealybug, Phenacoccus
rehi in Bangladesh. Bangladesh J Zool 9(1):1726 solani (Homoptera: Pseudococcidae) at three constant
Ali M (1995) Bionomics of sugarcane mealybug, temperatures. Appl Entomol Zool 41:573575
Kiritshenkella sacchari (Green) (Homoptera, Nikam ND, Patel BH, Korat DM (2010) Biology of inva-
Pseudococcidae). Pakistan J Zool 27(1):1519 sive mealy bug, Phenacoccus solenopsis Tinsley
Awadallah KT, Ammar ED, Tawk MFS, Rashad A (1979) (Hemiptera: Pseudococcidae) on cotton. Karnataka
Life-history of the white mealy-bug Ferrisia virgata J Agric Sci 23(4):649651
(Ckll.) (Homoptera, Pseudococcidae). Deutsche Pegoraro RA, Bellotti AC (1994) Biological aspects of
Entomologische Zeitschrift 26(1/3):101110 Pseudococcus mandio Williams (Homoptera:
Ayyar TVR (1939) The rice mealybug in South India. Pseudococcidae) in cassava [Portuguese]. Anais da
J Mysore Agric Exp Union 17:179188 Sociedade Entomologica do Brasil 23(2):203207
106 M. Mani and C. Shivaraju
Pramanik A, Ghose SK (1993) Experiments on the biol- Sahoo AK, Ghosh AB, Mandal SK, Maiti DK (1999)
ogy of the brinjal mealybug, Coccidohystrix insolita Study on the biology of the mealybug, Planococcus
(green) (Pseudococcidae: Homoptera). Ann Entomol minor (Maskell) Pseudococcidae: Hemiptera).
11(1):1518 J Interacademicia 3(1):4148
Rajendra A (1974) The biology and control of Vennila S, Deshmukh AJ, Pinjarkar D, Agarwal M,
Saccharicoccus sacchari Ckll. (Hom: Pseudococcidae) Ramamurthy VV, Joshi S, Kranthi KR, Bambawale
the pink mealy bug of sugar cane in Sri Lanka. Ceylon OM (2010) Biology of mealybug, Phenacoccus
J Biol Sci 11(1):2328 solenopsis on cotton in Central India. J Insect Sci
Razak TA, Ananthi DV, Jayanthi R (1994) Biology of the 10:119
sugarcane mealybug, Dysmicoccus carens Williams Walton VM, Pringle KL (2005) Developmental biology of
(Homoptera: Pseudococcidae). J Entomol Res vine mealybug, Planococcus cus (Signoret)
18(2):169174 (Homoptera: Pseudococcidae), and its parasitoid
Saha A, Ghosh A (2001) Biological studies on the mealy- Coccidoxenoides perminutus (Timberlake)
bug, Nipaecoccus viridis (Newstead) on various host (Hymenoptera: Encyrtidae). African Entomol
plants. Uttar Pradesh J Zool 21(1):7578 13(1):143147
Culturing of Mealybugs
7
M. Mani and C. Shivaraju
Culturing of mealybugs is essential to rear the laboratory in different countries on potato tubers
natural enemies, particularly parasitoids and and cucurbit fruits for the purpose of mass breed-
predators for use in the eld. Biological control ing of their parasites and predators (Ahmad and
programmes of mealybug species have relied on Ghani 1970). Large-scale multiplication of the
sprouting potatoes, pumpkins and butternut for mealybug for mass production of natural enemies
rearing of both mealybugs and their natural ene- was done on potato sprouts and pumpkin (Ahmad
mies. The ability to mass-rear the mealybugs is a and Ghani 1970. The rearing of Ferrisia virgata
vital step towards the culturing and colonisation (Cockrell) on brinjal has been reported by Rawat
of its natural enemies. An inexpensive mass- and Modi in India.
rearing technique and a nutritionally efcient but
simple diet have to be developed for the mealy-
bugs. The nutritional regime should be capable of 7.1 Potato Sprouts
producing and supporting great numbers of
mealybugs at low cost. An important requirement The use of potato sprouts as an insectary host for
for mass-rearing substrate is a long shelf life, the mealybugs culture was discovered by H. S.
which obviates the regular provision of fresh Smith and E. J. Branigan of the State insectary of
food. In this regard, butternut, pumpkins and USA (Branigan 1916). It was later modied by
sprout in potatoes have been found as suitable Smith and Armitage (1920), who found that
substrates for the mass-rearing of mealybugs white sprouts which developed in subdued light
(Johnson and Giliomee 2011). and at temperatures of 21.122.2 C were highly
In the large-scale production of mealybugs, acceptable to the mealybugs (Fig. 7.1).
potato sprouts or ripe pumpkins have been used The production method of P. citri on potato
in several countries. For mass rearing, a pure cul- sprouts is described in detail, as given by Fisher
ture of Planococcus citrii (Risso) must be main- (1963). The variety Red Bliss Triumph is pre-
tained in an isolated room solely for infesting ferred to Bounty or White Rose because it pro-
work (Finney and Fisher 1964). Species of duces sturdy sprouts highly acceptable to the
mealybugs have been satisfactorily reared in the mealybug and its natural enemies.
Potatoes after harvest should be stored for
more than 3 months at 2.2 C. Fans have to be
M. Mani (*) C. Shivaraju
provided to facilitate the circulation of air, which
Indian Institute of Horticultural Research,
Bangalore 560089, India reduces the temperature uctuation throughout
e-mail: mmani1949@yahoo.co.in the storeroom, and also reduces the decay
fostered by humidity. Planting trays are made of mum (properly bleached) potato sprouts.
redwood and their outside dimensions are Continuous weak light causes sprouts to become
18 4. Soil used is sandy silt, obtained from excessively etiolated and too much light causes
riverbed. Trays and soil can be sterilised prior to excessive leaf growth. Excessive long sprouts are
re-use. Approximately 3 months after harvest or pruned to 12 lengths. The time from planting
when sprouts begin to appear, the tubers are until infesting with mealybugs is usually 21 days
ready for planting. Medium to large-size sound in summer and 30 days in winter and in early
whole potatoes are used, and 2526 tubers are spring (Fig. 7.2).
placed about apart and on a layer of soil in Crawlers are removed from producing trays
the tray. They are covered with slightly damp- by allowing them to crawl into freshly cut short
ened soil. Immediately after planting, trays are leafy terminals of Switches of Pittosporus
placed on the racks of a thoroughly cleaned room undulatum placed among the sprouts. Schinus
and watered. Trays should be lled with brimful molle (California pepper) is also used in autumn.
of water every 4 or 5 days and watering should be Approximately 6 h after placing them on the
discontinued after the sprouts have been infested. trays of mealybugs, the switches are removed
Temperatures of 21.123.3 C appear to be opti- from the food room and placed on the fresh
mum for facilitating sprout growth and relative sprouts in the production from where the mealy-
humidity of 6084 % have given good results, bug crawlers move on to the sprouts as the
provided there is proper airow through the cul- switches dry out. During the transfer periods,
ture room. Control of light intensity in order to trays are not watered; light intensity is increased
minimise leaf growth and chlorophyll develop- and temperature is adjusted to 26.6 C. Another
ment is a critical factor in the production of opti- method is to remove every third tray in the row
7 Culturing of Mealybugs 109
and replace it with a tray containing long sprouts preferred host for the mass rearing of Oleander
that can be carefully bent over to interlace with mealybug Paracoccus burnerae (Brain) (Johnson
the over-producing sprouts of adjoining trays. and Giliomee 2011).
Stock from one mealybug tray will infest from 20 In India, the rearing of mealybugs on potato
to 25 trays. The optimum temperature for con- sprouts has been standardised by Joshi. The fol-
tinuous culture of mealybugs lies between 20 and lowing are the steps involved: (1) Procurement of
23.8 C (Fisher 1963). Sprouting potatoes are the medium-sized potatoes with well-developed
110 M. Mani and C. Shivaraju
eyes. (2) Cleaning with tap water and keeping in Cucurbita maxima Gil. (Mineo 1967). The prop-
a basin lled with sterilised sand. (3) Covering agation of citrus mealybugs on ripe pumpkins, C.
potatoes with sand and water moderately. (4) moschata D., has been outlined by Chacko et al.
Within a week or two, the sprouts grow to a (1978) and Singh (1978). The pumpkins with
length of 34 in.. (5) Removing the potatoes from ridges and grooves and a small stalk are selected,
sand and washing them with tap water. (6) which makes the handling easy. To prevent rot-
Keeping them in sunlight for a day so that the ting, the pumpkins are treated with 0.1 % beno-
sprouts turn green. (7) Infesting one sprout with myl (1 g/L). The wounds, if any on the pumpkin,
ve gravid females of mealybug. (8) Keeping are plugged with melted parafn wax. Ovisacs on
infested potatoes at 27 C and 5060 % relative P. citri /M. hirsutus are distributed over the pump-
humidity (R.H.). kin fruits or crawlers are dusted on the fruits. In
due course, the crawlers settle on all sides of the
pumpkin. The infested pumpkins are kept on
7.2 Pumpkin small tripod stands, which can be arranged in the
racks. In about a month, the mealybug population
The rearing of P. citri on cucurbits in USSR was covers the whole pumpkin. A temperature of 25
rst reported by Sysoev (1953). In Sicily (Italy), C is to be maintained in the rearing room
the mealybug P. citri was reared on pumpkin, (Chacko et al. 1978; Singh 1978) (Fig. 7.3).
The mealybugs are cultured on host plants. Initial Attempts were also made to rear the mealybugs
culturing is always done on natural host plants. on articial diet. Rearing the cassava mealybug,
Sometimes mass culturing is also done on the Phenacoccus manihoti, was done on a dened
host plants. A mass production protocol is avail- diet (Calatayud et al. 1998).
able for the production of papaya mealybug
Paracoccus marginatus Williams and Granara de
Willink on potato sprouts. However, it was very References
difcult to maintain potato plants in high-
temperature areas where spoilage of potatoes is a Ahmad R, Ghani MA (1970) A note on the rearing of
scale insects and mealybugs on potato tubers and
concern. Alternatively, a mass production tech-
cucurbit fruits. Tech Bull Commonw Inst Biol Control
nique has been developed using Hibiscus canna- 13:105107
binus for mass production of the papaya Branigan EJ (1916) A satisfactory method of rearing of
mealybug. Seeds were procured locally. Before mealybugs for use in parasite work. Calif State Hort
Comm Monthly Bull 5:304306
sowing, the seeds were spread on paper and
Calatayud PA, Delobel B, Guillaud J, Rahb Y (1998)
allowed to dry under sunlight for 13 h. Trays of Rearing the cassava mealybug, Phenacoccus manihoti
45 cm 30 cm 13 cm (l w h) with four to six on a dened diet. Entomol Exp Appl 86(3):325329
holes in the bottom to drain out excess water Chacko MJ, Bhat PK, Rao LVA, Deepak Singh MB,
Ramanarayan BP, Sreedharan K (1978) The use of the
were lled with a mixture of sand, soil and farm-
lady bird beetle, Cryptolaemus montrouzieri, for the
yard manure (1:1:1) up to 9 cm and H. cannabi- control of coffee mealybugs. J Coff Res 9:1419
nus seeds were sown for culturing the plants Finney CL, Fisher TW (1964) Culture of entomophagus
under laboratory conditions. The seeds started to insects and their hosts. In: DeBach PD (ed) Biological
control of insect pests and weeds. Chapman and Hall,
sprout within 2 days and attained a height of
London, pp 328353
40 cm within 2025 days, which is the suitable Fisher TW (1963) Mass culture of Cryptolaemus and
stage for infestation with papaya mealybug for Leptomastix, natural enemies of citrus mealybugs.
mass production purposes. Paracoccus margin- Bull Calif Agric Exp 797:39
Helen M, Chikkanna S, Balasaraswathi J, Ravikumar N,
atus, placed on 2025-day-old seedlings, was
Sakthivel N, Bindroo BB (2013) Hibiscus cannabinus
allowed to develop to the second instar, at which L. (Malvaceae) as an alternate host plant for mass-
time the plant was cut and transferred to plastic rearing of Paracoccus marginatus Williams and
containers having one thin layer of absorbent cot- Granara de Willink (Hemiptera: Pseudococcidae) and
their exotic parasitoids. In: Proceedings of 13th work-
ton covered with one layer of tissue paper. Based
shop of the IOBC Global working group on mass rear-
on the number of 2nd-instar mealybugs present, ing and quality assurance, Bangalore, India, 68 Nov
parasitoids were released at a ratio of 2:1 (para- 2013, pp 3940
sitoid to mealybug) into each container after Johnson T, Giliomee JH (2011) Evaluation of citrus, but-
ternut and sprouting potatoes as mass rearing sub-
which the cage was covered with muslin cloth.
strates for oleander mealybug, Paracoccus burnerae
For production of 20,000 parasitoids on the (Brain) (Hemiptera: Pseudococcidae). Afr J Biotechnol
mealybug using potatoes, the approximate cost 10:83208344
was Rs. 8700, whereas culturing done on H. can- Mineo G (1967) Cryptolaemus montrouzieri. Observations
on morphology and bionomics. Bull Inst Ent Agric
nabinus would require about Rs. 6700. It is con-
Oss Fitopat Palermo 6:99143
cluded that culturing of papaya mealybugs on H. Singh SP (1978) Propagation of a coccinellid beetle for
cannabinus is easy, economical and suitable for the biological control of citrus and coffee mealybugs.
tropical conditions and allows more effective Scientic Conference, CPA, Dec 1978, 2 p
Smith HS, Armitage HM (1920) Biological control of
mass production of the mealybug and its parasit-
mealybugs attacking citrus. Calif Agric Exp Stn Bull
oids (Helen et al. 2013). 9:104158
Sysoev AT (1953) The possibility of combining biological
and chemical methods in the control of pests of agri-
cultural crops. Dok Vseseovuz Akad Selkhoz Nauk
Lenin 18:2631
Mode of Spread of Mealybugs
8
M. Mani and C. Shivaraju
Root mealybugs spread through irrigation water Among the arthropods, ants have also been
from one spot to another. Besides, ood irriga- reported to disperse many mealybug species.
tion carries fallen leaves and other debris infested Ants are likely to carry the young mealybugs
with mealybugs from one spot to another. called as crawlers. Ants are known to transport
the mealybugs from plant to plant, between and
within elds, thus facilitating mealybug disper-
8.5 Air Currents sal. In California, it is often possible to see ant
Camponotus actually carrying the mealybugs
Some of the crawlers may be dispersed over lon- from its host plant, directly into the ants nest.
ger distances by air currents. Sticky trap collec- Ants are the primary or sole means of mealybug
tions revealed that Dysmicoccus neobrevipes dispersal in pineapple. Pheidole megacephala
Beardsley and D. brevipes (Cockerell) are dis- (F.) are seen carrying mealybugs from one pine-
persed by wind. In India, Paracoccus marginatus apple plant to another. The big-headed ant (P.
had spread very fast by wind dispersal of crawl- megacephala.), Argentine ant (Linepithema
ers from the state of Tamil Nadu to others. Aerial humile (Mayr)) and re ant (Solenopsis geminata
dispersal could be important in the colonization (F.)) are commonly found in the Hawaiian pine-
of mealybugs of new areas. apple agroecosystem, where they tend the mealy-
bugs for honeydew. These ants, especially P.
megacephala, have been blamed for dispersing
8.6 Animals mealybugs.
instars. Adult males are winged and capable of reducing the abundance of M. hirsutus with the
weak ight, but were caught on only 2 of the natural enemies, it would appear that many such
76 days of trapping. It is unlikely that the winged avenues for dispersal became ineffective.
males use the wind to assist them in dispersing to Presently, the spread of M. hirsutus is largely
new locations. Numbers of instars found in the limited to the transfer of mealybug life stages on
traps were positively related (p < 0.05) to the plants that are moved or especially ovisacs or
wind speed and to the square of the daily maxi- adult females on equipment. The cotton mealy-
mum temperatures. bug Ph. solenopsis was observed in 2006 and has
spread like wildre covering entire India within a
short time. Initially, hardly any parasitism was
8.12 Availability of Host Plant reported, but the absence of the parasitoid like
Aenasius bombawalei (later reported) aided in
Most of the mealybug species like spread of the mealybugs.
Maconellicoccus hirsutus (Green), Paracoccus
marginatus, Planococcus citri (Risso),
Nipaecoccus viridis (Newstead), Ferrisia virgata 8.14 Phoretic Method
(Cockerell), Phenacoccus solenopsis Tinsley,
Pseudococcus longispinus and Planococcus Reproductive females of the ant Acropyga epen-
lilacinus are highly polyphagous and known to dana Snelling participating in the ights are
attack hundreds of host plants aiding the spread known to carry the mealybug Rhizoecus colom-
of the mealybugs easily within the country. biensis (Hambleton) between mandibles indicat-
However, oligophagous cassava mealybug ing vertical transfer of mealybugs with their ant
Phenacoccus manihotti rst reported in 1973 hosts. Mealybugs and other scaled insects are
from Congo had become established in the whole known to cling to other insects like locusts and
cassava belt area by 1986, mainly due to the get dispersed during swarming. In another pho-
availability of cassava plants cultivated contigu- retic method, eggs and nymphs of Maconellicoccus
ously over a vast area in Africa. hirsutus were being transported by nymphs and
adult females of another mealybug, Ferrisia vir-
gata (Cockerell), in India.
8.13 Absence of Natural Enemies
then it quickly spread to all major grape-growing countries, causing severe damage. Since its
areas. The mango mealybug Rastrococcus accidental introduction into the island of Grenada
invadens Williams was accidentally introduced in in 1994, Maconellicoccus hirsutus, native of
Africa in the early 1980s from South East Asia South Asia, has been inexorably spreading
into Ghana and later spread to most of the African throughout several Caribbean islands.
Damage
9
M. Mani and C. Shivaraju
Mealybugs throughout the world cause a variety amino acids and other nitrogen compounds
of economic problems. The most obvious dam- (Franco et al. 2000; Gullan and Martin 2003;
age is caused by the sucking habits of these Silva and Mexia 1999; Tonkyn and Whitcomb
insects. The damage caused by the mealybugs is 1987). Thus, except for sucrose hydrolysis,
linked to sap intake. Heavy infestations often food digestion is hardly necessary. However,
cause stunting or death of the plant host. At times, organic compounds in phloem sap need to be
mealybugs have toxins and act as vectors of cer- concentrated before they can be absorbed, and
tain viruses detrimental to plant life. this occurs in the lter chamber, a specialized
component of the digestive system, which
enables the direct passage of water from the
9.1 Feeding Process anterior midgut to the Malpighian tubules,
and Endosymbionts thereby concentrating food in the midgut (Terra
and Ferreira 2003). The residue of ingested
Mealybugs are phloem feeders. As they feed, phloem sap, after digestion and assimilation in
they produce a sugary excretion (honeydew) the insect gut, is released from the anus as a
that supports the growth of sooty mould. sugar-rich material, the honeydew. Up to 90 %
Mealybugs feed by inserting their stylets of the ingested sugars may be egested in this
through the plant tissue to suck up sap from way (Mittler and Douglas 2003).
either phloem or mesophyll, or both. Males ter- Mealybugs have an obligatory association
minate their feeding towards the end of the sec- with prokaryotic endosymbionts, probably
ond nymphal stage. Generally, stylet because of the suboptimal nutrition furnished by
penetration is accomplished by the secretion of phloem sap, which lacks essential nutrients.
solidied saliva that forms a sheath around the These endosymbionts are believed to be
stylets. Similarly to other members of the sub- important for the nitrogen and sterol require-
order Sternorrhyncha, which includes scale ments of their hosts and may play a role in resis-
insects, aphids, psyllids and whiteies, mealy- tance to microbial pathogens or in detoxication
bugs consume a diet containing mainly carbo- of plant secondary compounds (Baumann 2005;
hydrates as well as limited amounts of free Gullan and Kosztarab 1997). Within their body
cavities, mealybugs have a structure, the bacteri-
M. Mani (*) C. Shivaraju ome, which comprises specialized cells, the bac-
Indian Institute of Horticultural Research, teriocytes, which harbour the primary
Bangalore 560089, India endosymbionts, that is, the P-endosymbionts
e-mail: mmani1949@yahoo.co.in
(Kono et al. 2008; Thao et al. 2002). The in the eld. For example, the citrus mealybug has
P-endosymbionts have the unusual property of been found on plants from 70 botanical families,
containing prokaryotic secondary endosymbi- 60 % of which are characterized as non-woody
onts, the S-endosymbionts, within their cyto- plants, whereas on the international scale, this
plasm (von Dohlen et al. 2001). During male mealybug is a pest of subtropical and tropical
development, the bacteriome progressively crops, such as citrus, persimmon, banana and
degenerates in the prepupa and pupa, and became custard apple, or it damages various types of
almost unrecognizable in the adult male (Kono plant species in interior landscapes, greenhouses
et al. 2008). in particular. Another example of the apparent
contradiction between the long lists of host plants
and the narrow ranges of damaged crops is the
9.1.1 Host Plants case of Pseudococcus cryptus Hempel; although
this mealybug is known from 35 host plant fami-
The various species of plants and animals have lies (Ben-Dov 2006), in Israel it causes damage
characteristic climatic requirements for growth, only to citrus trees. Under low pressure of natural
survival and reproduction, requirements that enemies, for example, when they spread in new
limit their geographic distribution, abundance environments, mealybugs are observed on rela-
and interactions with other species (Gutierrez tively large numbers of host plants, in contrast
et al. 1993). Mealybugs feed on a variety of her- with the situation when there is effective biologi-
baceous and woody plants, including the angio- cal control.
sperm, gymnosperm and fern families. However,
most of the species with known hosts develop on 9.1.1.1 Direct Damage
herbaceous plants, especially grasses (Poaceae) Mealybugs are phloem feeders that use long,
and composites (Asteraceae) (Ben-Dov 2006; slender mouthparts to suck out plant uids. Most
Kosztarab and Kozr 1988). As expected, infor- of the mealybugs can feed on the trunk, stem,
mation on the host ranges of mealybugs is mainly leaves, owers or fruits, and some on roots.
derived from observations of species of economic However, differences in the amount of damage
importance. Most species are apparently oligoph- caused by each species are often related to those
agous or stenophagous or monophagous, and factors that determine population size (e.g., num-
some are polyphagous (Ben-Dov 2006; Kosztarab ber of annual generations and female fecundity),
and Kozr 1988). However, such a characteriza- preferred feeding locations and temperature tol-
tion is problematic. Most of the economically erances. As the mealybugs feed, they excrete
important species are known to be associated carbohydrate-rich honeydew, which can accumu-
with long lists of hosts, and their performance late on the leaves and in the grape clusters, espe-
varies widely, ranging from development of high cially in late summer and early fall. The mealybug
population density, which eventually would kill icks honeydew away from its location, but it
the host plant, to poor development that renders still accumulates on the plant. It has long been
the survival of the population for several genera- noted that honeydew serves as a substrate for the
tions questionable. Plant growth conditions may development of sooty mould fungi that can result
strongly affect the success of the population: in further plant damage. For table-fruit growers,
under irrigation and fertilization, plant species any live or dead mealybugs and the honeydew or
become favourable hosts of mealybugs, whereas sooty moulds will cause cosmetic damage to fruit
in different environments the performance is usu- cluster and reduce its marketability. In most dried
ally poor. During laboratory studies, many of the fruits, juice and wine grapes, the contamination
mealybug pest species could be easily reared on from a small mealybug population, and the resul-
alternative hosts, such as potato sprouts or tant honeydew droplets, will not cause economic
squashes, which are not colonized by mealybugs damage. Although honeydew can be dissolved by
9 Damage 119
light rain and will dry in warm temperatures, problem to pineapple. Perhaps one of the most
when mealybug populations are severe, honey- important of these diseases is swollen shoot of
dew can accumulate to form a hard, wax-like cacao, transmitted by several mealybug vectors,
layer that covers the infested plant. A copious including Planococcoides njalensis (Laing) and
amount of honeydew gives the bark of the plant a Ferrisia virgala (Cockerell). This virus causes
water-soaked appearance. excessive damage to cacao trees each year.
Feeding damage can result in defoliation and,
after repeated annual infestations, cause vine
death. There are morpho-histological changes in 9.2 The Origin of Mealybug Pest
the plants due to the infestation of mealybugs on Status
plants like ramie (Boehmeria nivea), mulberry
(Morus alba), roselle (Hibiscus sabdariffa var. Similarly to other insect pests, mealybugs have
altissima) and mesta (Hibiscus cannabinus) diverse origins, including endemics, immigrants
infested with nymphs of Maconellicoccus hirsu- and mutants (Kim 1993). An indigenous species
tus (Green). Morphologically, linear growth of may become a serious pest: when a susceptible
the stem and petiole was arrested and their thick- crop species is introduced into the area, follow-
ness was increased. The leaf lamina was mark- ing environmental disturbance or as a result of
edly reduced and distorted. Histologically, the stress conditions. Invasive mealybug species
cells were enlarged and suffered reduced ligni- may attain pest status as soon as they success-
cation. There was an increase in the number of fully colonize a new territory, and affect nega-
stomata, which varied in the different plants. tively crop yield, which may happen when they
Mealybug (Rastrococcus invadens Williams) encounter a susceptible host, either local or
infestation of fruits caused a signicant reduction exotic. Planococcus citri (Risso) is an intro-
in weight and size of mango fruits, and also ash duced pest in most citrus-growing areas of the
content, crude bre and reducing sugars in Sri globe. It may weaken young citrus saplings, but
Lanka (Tobih et al. 2002). barely affects the growth of fruit-bearing trees;
the damage is mainly due to fruit infestation.
9.1.1.2 Indirect Damage Two mealybug population trends were shown to
In most of the regions, the transmission of occur in citrus orchards in the Mediterranean
viruses, rather than mealybug feeding or contam- region (Franco et al. 2004): (1) outbreak dynam-
ination, is the primary concern. Several species ics, whereby the percentage of infested fruits
of mealybugs are vector-virus disease in crops (mainly by Pl. citri) typically increases expo-
like banana, blackpepper, grapevine, cocoa, pine- nentially, with maximal values higher than 30 %
apple, sugarcane etc. Severe infestations can being recorded during mid- to late summer, and
result in defoliation, cluster infestation and rot, as (2) non-outbreak dynamics, whereby the per-
shown for a Planococcus cus (Signoret) infesta- centage of infested fruits does not increase sig-
tion. There is a slight leaf chlorosis and phloem nicantly or, alternatively, exhibits only a small,
disruption. Grapevine leafroll virus infections relatively linear increase, with maximal values
impact the berry development and growth by lower than 30 %. Three major causes may lead
delaying budbreak, owering and berry matura- to mealybug outbreaks: (1) a recent invasion by
tion, including changes in colour, reduced sugar an exotic mealybug species, (2) the application
content and increased acidity in fruit juice. of non-selective pesticides, which disrupt the
Mealybug toxins are rather important in some biological balance, and (3) the effect of environ-
areas. The pineapple wilt in the Hawaiian Islands mental factors that might inuence the tritrophic
involves the pineapple mealybug, Dysmicoccus interactions among host-plant/mealybug/natural
brevipes (Cockerell), which is a serious economic enemies.
120 M. Mani and C. Shivaraju
These were subdivided by Franco et al. (2004) The mechanisms involved in these two types of
as follows: outbreaks were discussed by Hardin et al. (1995),
and studied by Franco et al. (2004) with regard to
1. Recent invasion by exotic mealybug species the mealybug pests of citrus. Environmental fac-
(a) Lack of control by natural enemies tors (Cause 3) may also directly and indirectly
2. Application of non-selective pesticides affect the tritrophic interactions that develop
(a) Mortality differences between pests and between mealybugs, their host plants and their
their natural enemies natural enemies, thereby initiating mealybug out-
(b) Indirect effects of pesticides on natural breaks. Several mechanisms may be involved.
enemies, for example, elimination of their Host-plant characteristics may favour or be detri-
prey mental to the development, reproduction and sur-
(c) Effects on predator and parasitoid host vival of mealybugs (Boavida and Neuenschwander
interactions 1995; Calatayud et al. 1994b; Leru and Tertuliano
(d) Trophobiosis positive indirect effects of 1993; Nassar 2007; Tertuliano et al. 1993;
pesticides on pests, mediated through Wysoki et al. 1977; Yang and Sadof 1995). The
changes in the host plant resistance mechanisms of the host plant may
(e) Hormoligosis positive direct effects of become involved in both the xation (antixeno-
pesticides on pests sis) and the development of the mealybug (anti-
(f) Effects of pesticides on insect behavior biosis) (Tertuliano et al. 1993). Tertuliano and
(g) Effects of pesticides on interspecic com- Leru (1992) concluded that the different levels of
petition among phytophagous species of resistance to the cassava mealybug, P. manhioti,
different taxa which were observed in different varieties of cas-
3. Effect of environmental factors (tritrophic inter- sava, were not associated with the concentrations
actions: host-plant/mealybug/natural enemy) of amino acids or sugars, with the ratios between
(a) Host-plant susceptibility and/or host- these concentrations, or with the compositions of
plant characteristics amino acids obtained from leaf extracts. The
(b) Water stress identication and assay of cyanogenic and phe-
(c) Nitrogen fertilization nolic compounds in the phloem sap of cassava
(d) Weather and the honeydew of the cassava mealybug were
(e) Mealybug defences, for example, carried out by Calatayud et al. (1994a). Yang and
encapsulation Sadof (1995) showed that variegation in Coleus
(f) Mealybug refuges from natural enemies blumei could increase the abundance of the citrus
Spatial refuge (cryptic behavior), for mealybug, P. citri. Sadof et al. (2003) found that
example, under the bark and on roots the life-history characteristics of P. citri on
Temporal refuge: ant interactions Coleus blumei were not correlated with total
Other factors that may affect natural ene- amino acids and sucrose contents in stem exu-
mies, for example, intraguild predation dates, but were correlated negatively with the
and interference, hyperparasitoids proportions of shikimic acid precursors and posi-
tively with those of other nonessential amino
Cause 1 is well documented with regard to acids. Host-plant characteristics can also inu-
mealybug outbreaks and is mainly driven by the ence the performance of the natural enemies of
combination of host susceptibility and absence of mealybugs (Serrano and Lapointe 2002; Souissi
natural enemies in the invaded region (Ben-Dov and Leru 1997; Yang and Sadof 1997). Water-
1994; Blumberg et al. 1999; Muniappan et al. stressed plants may favour the population
2006; Nakahira and Arakawa 2006; Roltsch et al. increases of mealybugs (Calatayud et al. 2002;
2006; Williams and Granara de Willink 1992). Gutierrez et al. 1993; Lunderstadt 1998).
The use of non-selective pesticides (Cause 2) Mealybug life-history parameters/damage
may lead to resurgence and secondary outbreaks. may also be inuenced by the levels of nitrogen
9 Damage 121
fertilization and leaf nitrogen concentration; high Blumberg D, Ben-Dov Y, Mendel Z (1999) The citriculus
mealybug, Pseudococcus cryptus Hempel, and its
nitrogen concentrations were shown to lead to
natural enemies in Israel: history and present situation.
enhanced performance of the citrus mealybug, P. Entomologica Bari 33:233242
citri (Hogendorp et al. 2006). The antibiotic Boavida C, Neuenschwander P (1995) Inuence of host
resistance of two varieties of cassava mealybug plant on the mango mealybug, Rastrococcus invadens.
Entomol Exp Appl 76:179188
increased with the addition of nitrogen (Leru
Calatayud PA, Rahbe Y, Delobel B, Khuonghuu F,
et al. 1994). Survival of immature sugarcane Tertuliano M, Leru B (1994a) Inuence of secondary
mealybugs, S. sacchari, increased to a maximum compounds in the phloem sap of cassava on expres-
at a soluble nitrogen concentration of 320 mg L1 sion of antibiosis towards the mealybug Phenacoccus
manihoti. Entomol Exp Appl 72:4757
in sugarcane, and decreased at higher levels,
Calatayud PA, Tertuliano M, Leru B (1994b) Seasonal
whereas mealybug size increased with increasing changes in secondary compounds in the phloem sap of
nitrogen concentration over the whole tested cassava in relation to plant genotype and infestation by
range (Rae and Jones 1992). Weather conditions, Phenacoccus manihoti (Homoptera, Pseudococcidae).
Bull Entomol Res 84:453459
especially temperature and relative humidity, are
Calatayud PA, Polania MA, Seligmann CD, Bellotti AC
major ecological factors that affect both mealy- (2002) Inuence of water-stressed cassava on
bugs and their natural enemies (Chong and Phenacoccus herreni and three associated parasitoids.
Oetting 2007; Gutierrez et al. 1993, 2008a; Entomol Exp Appl 102:163175
Chong JH, Oetting RD (2007) Intraguild predation and
Nakahira and Arakawa 2006). Encapsulation
interference by the mealybug predator Cryptolalemus
may adversely affect the degree of biological montrouzieri on the parasitoid Leptomastix dactylopii.
control exerted by mealybug parasitoids, as it Biocontrol Sci Tech 17:933944
may either prevent the establishment of exotic Franco JC, Silva EB, Carvalho JP (2000) Mealybugs
(Hemiptera, Pseudococcidae) associated with citrus in
parasitoids in new regions or reduce parasitoid
Portugal. ISA Press, Lisbon (in Portuguese)
efcacy (Blumberg 1997). The cryptic behavior Franco JC, Suma P, da Silva EB, Blumberg D, Mendel Z
and tending of mealybugs by ants may, respec- (2004) Management strategies of mealybug pests of
tively, originate spatial and temporal refuges citrus in Mediterranean countries. Phytoparasitica
32:507522
from natural enemies. Several other factors may
Gullan PC, Kosztarab M (1997) Adaptations in scale
affect mealybugs natural enemies, which include insects. Annu Rev Entomol 42:2350
intraguild predation and interference (Chong and Gullan P, Martin JH (2003) Sternorrhyncha (jumping
Oetting 2007), and hyperparasitoids (Moore and plant lice, whiteies, aphids, and scale insects). In:
Resh VH, Card RT (eds) Encyclopedia of insects.
Cross 1992).
Academic, Amsterdam
Gutierrez AP, Neuenschwander P, Vanalphen JJM (1993)
Factors affecting biological control of cassava mealy-
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Mealybugs as Vectors
10
R. Selvarajan, V. Balasubramanian,
and B. Padmanaban
may continue to feed from the same sieve tube aphid transmitted viruses; apparently, the virus is
for several days. Xylem ingestion is also a pre- carried on or near the stylets of the mealybug.
dominant feeding behavior for some mealybug
species (Calatayud et al. 1994; Cid and Fereres
2010). 10.3 Plant Viruses Transmitted
by Mealybugs
Mealybugs are phloem feeders, and a minimum Nineteen species of mealybugs belonging to 13
inoculation time of 15 min is needed for success- genera are known to occur on Musaceae (Watson
ful transmission. The virus persists through the and Kubiriba 2005). The Banana streak virus
moult, and for 23 days in starved or feeding vec- (BSVs) (Fig. 10.1a) is transmitted by Planococcus
tors. All mealybug-transmitted viruses appear to citri (Risso) and Saccharicoccus sacchari
have a semi-persistent mode of transmission (Cockerell), both of which colonize bananas
based on retention times; however, the Grapevine (Lockhart et al. 1992). Sugarcane bacilliform
leafroll-associated virus 3 (GLRaV-3) was found virus (SCBV) is serologically related to BSVs
in the salivary glands of its mealybug vector, (Lockhart and Autrey 1988), and is reported to be
suggesting a circulative mode of transmission transmitted from sugarcane to banana by
(Cid et al. 2007). Mealybug-transmitted viruses Saccharicoccus sacchari (Cockerell) (Lockhart
appear to have a high rate of acquisition and a and Olszewski 1993). Experimental transmission
low rate of inoculation (Cid and Fereres 2010). of BSVs has also been demonstrated with the
Ants that tend to carry the mealybugs may move pink pineapple mealybug Dysmicoccus brevipes
them from one plant to another (Sether et al. (Cockerell) (Kubiriba et al. 2001) and
1998), and occasionally, long-distance dispersal Pseudococcus comstocki (Kuwana) (Su 1998).
by wind may also occur. An important factor Ferrisia virgata (striped mealybug) (Fig. 10.1b)
contributing to the slow rate of spread is that has been found to be able to transmit the Banana
newly infected trees are not infective for some streak Mysore Virus (BSMYV) from banana to
weeks, or even months, and the virus may not banana (Selvarajan et al. 2006). Meyer et al.
become fully systemic in large trees for at least 1 (2008) reported that the transmission of activated
year. Temperature-mediated mealybug activity episomal Banana streak OL virus (BSOLV) to
may be an important variable in transmission cv. Williams banana (Musa sp.) by three mealy
efciency, and the virus spread can occur through bug species, viz. Dysmicoccus brevipes,
the airborne dispersal of young, GLRaV-3- Planococcus citri and P. cus.
infected crawlers (Cabaleiro and Segura 1997). Planococcus citri transmitted episomal
Cacao swollen shoot virus (CSSV) is transmitted BSOLV and the Banana streak GF virus
in a semi-persistent mode, meaning that the virus (BSGFV) from tissue-culture derived plants of
is taken up into the vector's circulatory system FHIA-4 to cv. Williams plants. Using FHIA-TC
but does not replicate within it (Dzahini-Obiatey 10 as the donor plant for transmission, the vector
et al. 2010). The feeding period required for the transmitted a 100 % episomal BSOLV to
acquisition of the virus is a minimum of 20 min, Williamss plants after 3 months, and the num-
but optimally 24 days (Posnette and Robertson bers of mealybugs feeding on individual recipi-
1950). Once acquired, the virus can be transmit- ent plants during the inoculation access period
ted within 15 min, but optimal transmission (IAP) ranged from 2 to 25 (Meyer et al. 2008).
occurs 210 h after acquisition. No transmission Episomal BSVs were transmitted by D. brevipes.
of the virus occurs through the mealybug eggs. At 3 months post transmission, the virus was
The relationship between the CSSV and mealy- detected and symptoms had appeared. Due to the
bugs has some similarities to the non- persistent reluctance of the mealybugs to move to Musa spp.,
10 Mealybugs as Vectors 125
a b c
e f g
h i j
Fig. 10.1 (a) Electron micrograph of BSV; (b) Ferrisia Electron micrograph of GVB; (g) Vine mealybug; (h)
virgata feeding on banana; (c) Pink pineapple mealybug, Symptoms of BSV in banana; (i) GLRaV infected grape-
Dysmicoccus brevipes; (d) Gray pineapple mealybug, D. vine; (j) Symptoms of mealybug wilt of pineapple
neobrevipes; (e) Electron micrograph of GLRaV; (f)
the low numbers survived the inoculation access et al. 2008). The fact that none of the mealybug
period; the virus was transmitted from the TC-5 species were able to transmit integrated BSOLV
to the William banana. However, no episomal from the FHIA-4 to Williams proves that the inte-
BSGFV could be transmitted by D. brevipes from grated form of BSV is not likely to be transmitted
the FH-4 donor to the recipient plants (Meyer by mealybugs; even highly efcient mealybugs
126 R. Selvarajan et al.
such as P. citri were unable to transfer any inte- PMWaV-2 infection and mealybug feeding are
grated viral sequences to the receptor plants. necessary for the development of mealybug wilt
Episomal BSV in tissue culture-derived tetra- disease (Hu and Sether 1999a, b; Sether and Hu
ploids is highly transmissible by efcient mealy- 2002a, b). All pineapple plants with wilt disease
bug vectors to Cavendish varieties. have PMWaV-2 infections, but not necessarily
Cocoa swollen shoot virus (CSSV), a badna- PMWaV-1 infections (Hu et al. 1997; Sether and
virus, is transmitted by at least 14 species of Hu 2002a). Several species of ants are associated
mealybugs of the family Pseudococcidae within with mealybugs (Beardsley et al. 1982; Carter
the Coccoidae (Roivainen 1976), but 1963). These ants assist in the establishment of
Planococcoides njalensis and Planococcus citri mealybug colonies, consuming the honeydew
are the most important vectors (Dongo and produced by the mealybugs (Petty and Tustin
Orisajo 2007). Piper Yellow Mottle Virus 1993), and can have a suppressive effect on the
(PYMV) is transmitted by the citrus mealybug, natural enemies of mealybugs (Jahn 1992).
Planococcus citri (Lockhart et al. 1997). Bhat Sether et al. (1998) reported that presence of ants
et al. (2003) reported that PYMV could easily be was correlated with an increased rate of virus
transmitted by the mealybugs (Ferrisia virgata) spread when caged with D. brevipes. All stages
from naturally diseased black pepper to healthy of D. neobrevipes acquire PMWaV, although
seedlings of black pepper. The initial symptoms vector efciency decreased signicantly in older
of the disease, like vein clearing and chlorotic adult females; the probability of a single third-
mottle, could be seen in 14 of the 20 test plants in instar immature transmitting the virus was 0.04.
5 weeks after inoculation. Macanawai et al. Both the species of the mealybugs acquired and
(2005) reported that the Taro bacilliform virus transmitted the PMWaV from infected pineapple
(TaBV) is transmitted by Pseudococcus material.
solomonensis. The Grapevine leafroll disease is caused by
grapevine leafroll-associated viruses (GLRaVs)
(Fig. 10.1e). These viruses are common in vine-
10.4 Viruses Belonging yards worldwide, and are often associated with
to Closteroviridae vitiviruses that are involved in the rugose wood
complex of grapevines. Ten mealybug species are
Mealybug-vectored viruses often exist as a com- known as vectors of one or several of these grape-
plex of viruses, such as the mealybug wilt of vine viruses, including the apple mealybug
pineapple complex, which is made up of three Phenacoccus aceris, which is widespread, and is
pineapple mealybug wilt-associated viruses able to transmit the Grapevine leafroll-associated
(PMWaV) (Sether et al. 1998, 2005; Sether and virus-1 and -3 (GLRaV-1 and -3). Vitiviruses,
Hu 2002a, b) and Grapevine leafroll-associated namely Grapevine virus A (GVA), Grapevine
viruses. Mealybug wilt of pineapple is a major virus B (GVB) (Fig. 10.1f), Grapevine virus D
constraint in the global production of pineapple (GVD) and Grapevine virus E (GVD), infect
(Carter 1934, 1942; Rohrbach et al. 1988; grape vines, and these are transmitted by the
Wakman et al. 1995). Carter (1934, 1942, 1949, members of several insect genera (Pseudococcus,
1962) found an association between mealybugs, Planococcus, Phenacoccus, Heliococcus,
particularly the pink pineapple mealybug, Neopulvinaria, Parthenolecanium, Cavariella
Dysmicoccus brevipes (Cockerell), (Fig. 10.1c) and Ovatus) in a semi-persistent manner (La
and the gray pineapple mealybug, D. neobrevipes Notte et al. 1997; Rosciglione et al. 1983; Garau
(Beardsley) (Fig. 10.1d), and wilt throughout the et al. 1995). Tsai et al. (2010; Le Maguet et al.
pineapple-growing regions of the world. 2012) studied the virusvector specicity analy-
PMWaV-1 infections are correlated with growth sis for mealybug transmission of GLRaVs. Plants
reductions of the plant crop (Sether and Hu infected with several GLRaVs virus species were
1998), and yield reductions in the ratoon crop. screened for vector transmission by the mealybug
10 Mealybugs as Vectors 127
species Planococcus cus and Pseudococoucus (AAP) and peaked with a 24-h AAP, after which
longispinus. The results revealed that P. longispi- the transmission rate remained constant. In addi-
nus had transmitted the GLRaV-9 to the inocu- tion, the GLRaV-3 was found not to have been
lated plants, and showed that 18 % of the transovarially transmitted from infected females
inoculated plats were positive for GLRaV-9, but to their progeny (Table 10.1).
none of the inoculated plants were found positive Mealybugs are less mobile on the plant com-
for GLRaV-5, tested 9 months after the inocula- pared with groups of vectors such as aphids and
tion. Planococcus cus transmitted the GLRaV- leaf hoppers, a feature that makes them relatively
1,3,4,5,9 and GVA. This study showed that there inefcient as virus vectors. Mostly, the mealybug-
was no evidence of mealybugGLRaV specic- transmitted viruses appear to have a semi-
ity. Tsai et al. (2008) reported that the vine persistent mode of transmission based on
mealybug (Planococcus cus) (Fig. 10.1g) trans- retention times. Mealybug-transmitted viruses
mits GLRaV- 3 in a semi-persistent manner. First appear to have a high rate of acquisition and low
instars were more efcient vectors than adult rate of inoculation. Mealybug-vectored viruses
mealybugs, but the GLRaV-3 transmission lacked often exist as a complex of viruses, such as the
a latent period in the vector. Virus transmission mealybug-associated viruses. Badnaviruses such
occurred with a 1-h acquisition access period as PYMV, BSVs TaBV and CSSV have been
128 R. Selvarajan et al.
shown to transmit by different mealybug species. 34 years (Fig. 10.1j) (Crowdy and Posnette
In all of these studies, the interaction of the 1947). The impact of mealybug feeding and
mealybug vector with the virus and the host is Pineapple mealybug wilt associated virus-1
lacking; hence it is necessary to generate funda- (PMWaV-1), PMWaV-2 infection on pineapple
mental knowledge about the interaction of the yield and the spread of PMWaV-1 and mealybug
vector, the host and the virus system to develop wilt of pineapple (MWP) were evaluated under
effective disease management strategies for viral eld conditions; the results showed a 35 % reduc-
diseases. Epidemiological studies are also tion in yield when compared with PMWaV-free
required to predict the spread of the plant viruses plants (Sether and Hu 2002b). If MWP develops
through mealybugs, and the changing climatic during the rst 3 months of the plant crop, it can
conditions need to be considered while develop- lead to a 55 % reduction in average fruit weight,
ing forecasting models of disease spread. compared with fruits from PMWaV-free plants.
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Economic Importance
11
M. Mani and C. Shivaraju
Mealybugs are widely distributed phytophagous discoloured welts on the rind of the fruit,
insects, often with broad host ranges. There are ower, etc.
approximately 2000 described mealybug species 2. Mealybugs excrete copious quantities of hon-
worldwide. According to Millar et al. (2002), 158 eydew, which is a substrate for the fungus,
species of mealybugs are recognized as pests. sooty mould. Sooty mould is black in colour
Mealybug is a pest, which can have a consider- and may stain the fruit/ower decreasing the
able negative economic impact on a wide range packout percentage as well as causing a delay
of crops and ornamentals. In the last 30 years, in fruit colour development. Photosynthetic
there have been several major outbreaks of potential, especially of young trees, may be
mealybugs causing alarming damage to crops, as negatively affected if sooty mould infection is
a result of invasion/accidental introductions. severe.
Losses and costs of controlling mealybugs in 3. Mealybug is a phytosanitary pest in some
Georgia (USA) in 1996 were estimated at about export markets (USA, Japan) and if found on
$9.8 million. Damage and costs of controlling the fruit/ower destined for these markets can
pink hibiscus mealybug in the United States were result in rejection of the consignment and
recently estimated at $700 million annually. In could place these important markets at risk for
South Africa, costs for control of vine mealybug the future.
in vineyards were estimated at around $100 per 4. Mealybugs act as vectors of plant virus dis-
hectare per season. Most notorious mealybug ease causing heavy losses. Several mealybugs
species are polyphagous, and have become seri- are responsible for transmission of Grapevine-
ous pests of different crops under different leafroll-associated virus (GLRaV), and the
environments. virus infection was predicted to spread with
Economic damage can happen in four ways: the economic impact of Grapevine-leafroll-
associated virus-3 (GLRaV-3) infection
1. A high population of mealybug can lead to exceeding 10,000 dollars per ha, annually in
fruit, ower/leaf drop, fruit/ower deforma- South Africa.
tion (high shoulders) and development of
Mealybugs spread between continents through
M. Mani (*) C. Shivaraju
international trade. In the United States, there are
Indian Institute of Horticultural Research,
Bangalore 560089, India 350 species of mealybugs. Approximately 70 %
e-mail: mmani1949@yahoo.co.in of the 66 mealybug species that are considered as
pests are invasive. Invasive mealybugs in also known to transmit the virus known as cholo-
California are serious pests of several economi- rotic streak (Williams 2004). Birendracoccus
cally important crops. There are several other saccharifolii (Green) is a major pest of sugarcane
mealybugs that reveal the extent of damage and in India and a vector of spike disease (Ali 1962).
economic losses. In New Zealand, most of the Coccidohystrix insolita (Green) has been a seri-
known 114 species of mealybugs are found only ous pest of brinjal, egg plant/aubergine, in Bihar,
on native plants. Three cosmopolitan and inva- West Bengal, Tamil Nadu, Kerala and several
sive Pseudococcus species are frequently occur- other states in India (Williams 2004). Economic
ring pests of horticultural crops in the country, damage by mealybugs on brinjal was reported in
where they account for more than 99 % of the Pakistan and also in other Asian countries (Arif
mealybug fauna in orchards and vineyards et al. 2009).
(Charles 1993). In France, scale insects, includ- Dysmicoccus boninsis (Kuwana) is a wide-
ing mealybugs, represent 31 % (Streito and spread pest of sugarcane causing economic dam-
Martinez 2005) of the newly introduced species age (Ben-Dov 1994). Dysmicoccus brevipes
in recent years, although all mealybug pests on (Cockerell) is a well-known pest of pineapples
grapevine are native (Sforza 2008). Likewise, in worldwide and also coffee. It acts as vector of
many countries, there were serious economic pineapple wilt in Hawaii and several other coun-
losses caused by mealybugs. tries. It is one of the principal pests of mango in
Rhodesgrass mealybug, Antonina graminis Okinawa. D. brevipes was reported on oilpalm-
(Maskell), has been a major pest of many pasture infesting leaves, inorescence and ripe fruit
grasses and lawns, and to some extent on bam- bunches in India (Ponnamma 1999). Dysmicoccus
boos in various parts of the world. It had com- neobrevipes (Beardsley) is common in Hawaii
pletely destroyed thousands of acres of good and also in southern Asia. It has caused severe
pasture land. Injury is rst indicated by the stunt- loss to tube rose growers in India. Dysmicoccus
ing and reduction in the overall size of individual grassii (Leonardi) has been reported as a pest of
grass clumps, with darkening of the leaves and banana in Canary Islands (Beardsley 1964a, b)
eventual death of the host plant. Death of seed- and heavy infestations on plantain in Nigeria.
ling plants is known to occur in about 3 weeks. Ehrhornia cupressi (Ehrhorn) is a serious pest,
Conventional control is difcult because of the which caused the destruction of cypress hedges
position of the mealybug on its host. The success in California (Herbert 1920).
in controlling this mealybug in Texas is by the Gilli mealybug Ferrisia gilli (Gullan) is the
introduction of the parasitoid Neodusmetia sang- primary pest of pistachio covering over 3000
wani (Subba Rao) from India (Dean et al. 1979). acres of pistachios in California. It is also known
The cost of the control programme was estimated to attack and cause huge losses to a wide range of
at that time at $0.2 million, resulting in the sav- crops such as almonds, grapes, stone fruits.
ings of about US$200 dollars per annum. The striped mealybug Ferrisia virgata
Subsequently, colonies of the parasitoid have (Cockrell) has been of some concern to several
been sent elsewhere in the New World. Heavy countries. In the past few years, however, heavy
infestations of the mealybug Antonina pretiosa infestations were noticed on many ornamental
(Ferris) produce unsightly condition of the bam- plants. This has caused some alarm as this spe-
boo (McKenzie 1967). Brevennia rehi (Lindinger) cies is reported as an important pest, especially to
is an important pest of rice in India, Pakistan, cotton. It is found normally above ground on the
Burma, Indonesia, Bangladesh and some other foliage where it causes the usual honeydew-sooty
countries causing severe loss of the crop espe- mould-type damage. During severe weather con-
cially in the dry seasons. Grains from mealybug- ditions, in Africa at least, it may move to the
infested plants did not develop properly and that crown and roots of its host. This mealybug is
they tasted bitter, and if present in normal food, found on a wide range of hosts. It caused eco-
they spoilt the avour after being cooked. It is nomic losses to citrus, guava, custard apple,
11 Economic Importance 133
mango, cotton, pomegranate, pummelo, tuberose, to 100 % loss in grapevine and mulberry. It is also
pepper, jackfruit, poinsettia, Acalypha, known to attack other crop plants, guava, pome-
Caesalpinia, etc. in India (Mani and granate, custard apple, acid lime, Phalsa, hibis-
Krishnamoorthy 1993), as well as pepper in India cus, ber, sapota, okra, etc. in different countries.
and jute in Bangladesh. Formicococcus robustus Introduced natural enemies, mainly the parasit-
(Ezzat and McConnell) is only known from the oid Anagyrus kamali (Moursi) (already known in
Indian region but it is sometimes intercepted at the Old World) and Gyranusoidea indica (Shafee,
port inspection elsewhere. Although a polypha- Alam and Agarwal) and the predator
gous species, it is frequently found on mango, Cryptolaemus montrouzieri (Mulsant), have
and in Pakistan it is reported as a serious pest. brought the mealybug under control in Egypt,
Kiritshenkella sacchari (Green) is known to West Indies, the United States, etc.
cause severe loss to sugarcane growers in India. Maconelicoccus hirsutus was detected in teak
Another example that indicates the high eco- plantations in 2004 in the Banderas valley in
nomic importance of a polyphagous mealybug is Mexico. A biological control programme was
the pink hibiscus mealybug, Maconellicoccus initiated in May 2004 to release 210,000 of the
hirsutus (Green). This mealybug is indigenous to predator Cryptolaemus montrouzieri on 150 ha of
southern Asia, and actually is considered a poten- land. Damage to trees was reduced by 92 % (Villa
tially serious pest in the United States, because of Castillo 2006). Mizococcus sacchari (Takshashi)
its extremely broad range of economically impor- was very injurious to sugarcane in Taiwan
tant hosts, including citrus, ornamentals, vegeta- (Takahashi 1928). Its presence in European and
bles and the native American ora. It was rst Mediterranean Plant Protection Organization
reported in the Western Hemisphere in Hawaii in (EPPO) countries would probably affect export
1984, and later in Grenada in 1994; subsequently markets, since it is regulated as a quarantine pest
it has spread rapidly through the Caribbean by many countries in other continents.
islands and to southern California (1999) and Nipaecoccus viridis (Maskell) is widespread
Florida (2002). Without control, the economic throughout tropics and subtropics causing eco-
impact of M. hirsutus to U.S. agriculture has nomic losses to numerous crop plants including
been estimated at $750 million per year (Hall citrus, pomegranate, guava, grapes, ber, jackfruit,
et al. 2008). The same pink hibiscus mealybug mango, custard apple and pummelo in several
was introduced accidentally to the Caribbean counties. In India, it is a pest of stored potatoes.
area in 199394, and has since spread beyond, Cotton is often attacked, when gall-like swellings
eventually reaching the United States. This dam- appear on terminal shoots, and tea is often heav-
aging species was rapidly identied by taxono- ily infested. On Artocarpus spp., large aggrega-
mists such as Maconellicoccus hirsutus (Green); tions of the mealybug lead to drying of the shoots.
its biological control was described in detail by In South Africa, N. viridis is a major pest of cit-
Kairo et al. (2000), with discussion of the costs rus, and in Okinawa it is one of the principal pests
and benets. M. hirsutus is widely distributed of mango. When rst introduced into Jordan in
throughout southern Asia, Africa and other parts 1993, apparently without natural enemies, infes-
of the Old World including Australia. M. hirsutus tations sometimes resulted in total loss of the cit-
is reported as a vector of virus disease on cocoa rus crop. In Egypt, a severe outbreak of N. viridis
in Zanzibar and other plants in East Africa, which occurred on lebbak trees. Niapecoccus nipae
causes growth arrest and branch distortion (De (Makell) has become serious pest of avocado and
Lotto 1967). Maconellicoccus hirsutus gets guava in Hawaii (Zimmerman 1948) and Puerto
ranked as one of the most important polyphagous Rico (Martorell 1940). The species is now con-
mealybugs in southern Asia, especially in India trolled successfully in Hawaii by the parasitoid
still causing damage in parts of India (Mani et al. Pseudophycus uitilis (Timberlake). Nipaecoccus
2011). The pink hibiscus mealybug is a major nipae was also known to cause serious damage to
pest of grapevine in peninsular India causing up coconut in Bermuda (Bennet and Hughes 1959).
134 M. Mani and C. Shivaraju
Palmicultor palmarum (Ehrhorn) is reported calyxes of apple and pears grown for export has
to infest 5 % of coconut palms in Bangladesh and caused concern in Australia and New Zealand
India. The species attacks the spear leaves of oil (Ward 1966). Phenacoccus madeirensis (Green)
palm in India. It sometimes occurs deep in the is a common polyphagous mealybug in much of
brous material covering palm stems, where it is the New World, Africa and the Pacic region.
difcult for chemical insecticides to penetrate. This mealybug is injurious to potatoes (Solanum
Paracoccus marginatus (Williams and Granara tuberosum) in Peru, and the growth of associ-
de Willink) causes serious economic losses to the ated sooty moulds causes malformation and
tune of several crores of rupees to more than 90 damage to leaves of other plants in Japan,
plant species particularly to the papaya, tapioca where it has been reported recently. It has
and mulberry damage in more than 53 countries invaded India recently and found to be severe
including India. Paracoccus marginatus has on tapioca.
become a serious pest in the Caribbean islands, Phenacoccus manihoti (Matile-Ferrero)
where it attacks numerous plant species, espe- appeared on cassava in Africa in 1973, and soon
cially papaya. The mealybug has now reached the spread throughout the whole cassava belt. The
southern United States. The mealybug reached introduction of the parasitoid Apoanagyrus lopezi
Guam and Palau on C. papaya; these islands are (De Santis) from South America to Africa and
possible sources for future incursions into the the success of the biological control programme
Pacic area and southern Asia. Biological control against P. manihoti had been well documented by
of Pa. marginatus with Acerophagus papayae Herren and Neuenschwander (1991). The tre-
(Noyes and Schauff) saved the silk, papaya and mendous success is credited with preventing the
tapioca industry from the loss worth to 2000 malnutrition of millions of Africans and may
crores rupees in India alone (Mani and Shivaraju well be the most important example of classical
2012). Similar economic benets were realized biological control ever. Zeddies et al. (2001) cal-
in several other countries. Hatting (1993) reported culated the total costs and benets of this biologi-
Paracoccus burnerae (Brain) as the most impor- cal control programme for 27 African countries
tant pest on citrus in South Africa. over a 40-year period (19742013) under differ-
Phenacoccus aceris (Signoret) has become a ent scenarios, such as transport, loss of crop and
serious threat to apple, pear, plum and other fruit even the price of maize as a possible substitute.
trees in Miane, British Columbia, Nova Scotia, Based on the total cost of biological control at
California and South Africa. The parasitoid US$ 47 million, the benets from different sce-
Allotropa utilis (Muesbeck) was introduced to narios range mainly from 199:1 (or US$ 9.4 bil-
British Columbia where it became well estab- lion) to 430:1 (or US$ 202 billion). Although the
lished. This was considered one of the outstand- initial cost of identication of the mealybug was
ing successes of classical biological control. negligible, there was a taxonomic advantage in
Phenacoccus gossypii (Townsend and Cockerell) that the costs included funds set aside for a study
is widely distributed in many countries. It is a of the mealybugs of Central and South America
pest of numerous owering plants in nurseries by Williams and Granara de Willink (1992).
and greenhouses, and in natural environments. Phenacoccus manihoti remains a threat to the
This mealybug, which is most often found on the cassava in the areas of southern Asia, as does the
foliage of its host, apparently causes as much yellow cassava mealybug, P. herreni, which still
damage to its host plants. Phenacoccus solani causes problems in South America. Reduction of
(Ferris) has probably been introduced recently P. herreni populations is under way, mainly
and is now established in southern Asia. There through the introduction of the parasitoids
are reports that it is a pest of stored potatoes in Apoanagyrus diversicomis (Howard) and
North America, and heavy infestations have been Acerophagus coccois (Smith) (Bento et al. 1999).
found on tobacco in Zimbabwe. Presence of The most trenchant point concerning the parthe-
Phenacoccus graminicola (Leonardi) under the nogenetic species P. manihoti is that an outbreak
11 Economic Importance 135
could occur in southern Asia with the accidental Planococcus citri (Risso) is one of the most
introduction of just a single immature specimen. cosmopolitan mealybugs. It is considered a seri-
Phenacoccus solani (Ferris) and Ph. solenop- ous pest of citrus in many parts of the world dam-
sis (Tinsley) are examples of invasive pests of aging many other eld crops in tropics and
annual crops; they cause heavy damage to green subtropics as well as greenhouse in temperate
pepper in Israel and cotton in the Indian subcon- regions. The mealybug is known to attack mainly
tinent (Ben-Dov 2005; Hodgson et al. 2008; subtropical fruit trees and also olive under
Nakahira and Arakawa 2006). The damage Mediterranean climate conditions and ornamen-
caused by mealybugs is linked to sap uptake, tal plants in interior landscapes in cooler zones
honeydew secretion and associated sooty mould (Ben-Dov 1994; Franco et al. 2004). The citrus
development, toxin injection and virus transmis- mealybug has become a key pest in the mint and
sion, although the presence of the insects may tarragon industry in Israel. This cosmopolitan
itself lead to economic losses (Franco et al. 2000; species was probably the rst recorded as pest in
McKenzie 1967; Panis 1969). The cotton mealy- southern Asia. Chemical control of this insect is
bug, Phenacoccus solenopsis, native of the amazingly difcult. Planococcus citri is known
United States (New Mexico) invaded several to cause up to 38 65 % damage on various citrus
countries Central America, the Caribbean and species (sweet orange, acid lime and lemon),
Ecuador (Argentina, Brazil, Ghana, Colombia, pummelo, guava, grapes (60 % loss), ber, sapota,
Nigeria, Asia (Pakistan, India and China)). It is a pomegranate, custard apple, crossandra, coffee,
major pest posing a severe threat to the cotton etc. in India (Manjunath 1986; Mani 2001).
crop in India and vegetable growing areas of Biological control of P. citri with natural enemies
Thailand and several ornamental plants in many saved several citrus orchards in India, the United
countries. This mealybug caused economic dam- States, Italy, Australia and South Africa.
age in India and Pakistan reducing the yields up Planococcus is also listed as a vector of Ceylon
to 450 %. Phenacoccus solenopsis caused a loss cocoa virus in Sri Lanka and also Grapevine virus
of several lakhs of rupees to cotton growers in (GVA). Planococcus cus (Signoret) is a pest of
India alone. Phenacoccus gossypii (Townsend grapevine in the Mediterranean region, South
and Cockerell) is widely distributed in many Africa, Pakistan, Argentina, Georgia and
countries. It is a pest of numerous owering California, causing heavy losses to grape grow-
plants in nurseries and greenhouses, and in natu- ers. It also transmits the grapevine leafroll virus.
ral environments. This mealybug, which is most Planococcus kenyae (Le Pelley) is popularly
often found on the foliage of its host, apparently known as coffee mealybug. It has caused heavy
causes as much damage to its host as the citrus losses to coffee growers in Uganda, Tanzania and
mealybug, Planococcus citri (Risso). Heavy Kenya (Bigger 2009).
infestations of Ph. solani have been recorded on Planococcus lilacinus (Cockerell) is one of
tobacco in Zimbabwe. the most common in southern Asia and reports of
Phenacoccus saccharifolii (Green) is known damage vary. It is known to attack and cause seri-
to attack sugarcane in India, Nepal and Pakistan. ous economic losses to cocoa, guava, ber, citrus,
In Bihar (India), infestation causes leaves and black pepper, cashew, pomegranate, guava,
internodes to become drastically reduced so that sapota, coffee, chow chow, mango, etc. (Mani
the cane can resemble a spike. Young sugarcane 2001). Planococcus lilacinus is known to trans-
plants have been severely damaged by this spe- mit Ceylon cocoa virus in parts of Sri Lanka.
cies in West Bengal (India). Planococcoides Planococcus cus is a serious pest of grapevine
nijalensis (Laing) is the dominant vector of the in the Mediterranean region, South Africa,
cocoa swollen shoot virus in African countries. It Argentina, Georgia and Pakistan. It is also found
is also known to attack cashew, Annona, silk cot- transmitting GVA and grape leafroll virus (Ben-
ton, pineapple, Acacia, Albizia, Caesalpinia, Dov 1994; Daane et al. 2006; Zada et al. 2008).
Erythrina, coffee, Clerodendron, etc. Planococcus minor (Maskell) is a common
136 M. Mani and C. Shivaraju
species on economically important plats particu- a pest in greenhouses and nurseries, but is also
larly cocoa throughout its geographical range. found out of doors in warmer areas. It has been
Trees (Cupressus and Juniferus) infested with reported as a pest of avocados, grapes and citrus.
Planococcus ovae (Nasonov) suffer from dieback Severe infestations have been reported on black
of twigs, heavy accumulation of honey dew and pepper in India. The mealybug is a target pest for
decline of trees in Italy (Ben-Dov 1994). classical biological control in Australia, and the
Planococcus kraunhiae (Kuwana) is widely species has caused damage to avocados in Israel
spread in California, Taiwan, China, Japan dam- in recent years. It also acts as a vector of grape
aging fruit tress such as pears, grapes, persim- leafroll virus. The orchid mealybug Pseudococcus
mons, banana, citrus, gs, etc. Planococcus microcirculus (McKenzie) has caused many
minor Maskell is a common species of many eco- problems to orchid growers (McKenzie 1967). It
nomically important plants including cocoa. is found primarily on the roots of its host but
Planococcus ovae (Nasanov) is widely distrib- crawls to the foliage and leaf sheaths when infes-
uted in Neotropical and Palaearctic regions on tations become heavy. The type of damage is the
Anthurium, Cupress and Juniperus trees. Infested normal form of unsightly contamination with the
trees suffer from dieback. production of honeydew during heavy infesta-
Several members of the genus Pseudococcus, tions. Pseudococcus maritimus (Ehrhorn) is
for example, Ps. calceolariae (Maskell), Ps. lon- another species important primarily to grapes and
gispinus (Targioni-Tozzetti) and Ps. viburni pears in some countries. The presence of these
(Signoret), are important pests of apple, pear and mealybugs on the ripe marketed grapes results in
vineyards in New Zealand (Charles 1993), serious economic loss to the growers. Heavy
whereas around the Mediterranean they are con- infestations cause the grapes to crack, allowing
sidered mainly as pests of citrus, persimmon and mould contamination. Pseudococcus viburmi
several other subtropical fruits (Franco et al. (Signoret) is most common in tropical and tem-
2004). Pseudococcus comstocki (Kuwana) is a perate areas but it is not widespread in southern
serious pest on apple, mulberry, pears, peach in Asia. It may have been overlooked, however,
the United States and Japan. The citriculus owing to its cryptic habit of living on roots. In
mealybug, Pseudococcus cryptus (Hempel), is a Australia, it causes damage to lawns and tubers,
major pest of citrus in the east Mediterranean and is a target species there for classical biologi-
region, and it attacks coffee roots in Asia and cal control. It is causing damage to Californias
South America (Ben-Dov 1994; Williams and coastal vineyards.
Granara de Willink 1992). It is widespread and a Following the introduction of the cassava
polyphagous mealybug species and appears to be mealybug into Africa, another introduced mealy-
kept under control by natural enemies in southern bug Rastrococcus invadens (Williams) appeared
Asia. Citrus and coconut are its favourite host in West Africa in 198182, causing extensive
plants, and infestations are known to occur on oil damage to fruit trees including mango (Williams
palm in India. It is widely distributed in Southeast 1986b). This mealybug was already known from
Asia, Tropical Africa, Middle East Meditrranean India and Pakistan (Narasimham and Chako
and South America. It is particularly a pest of cit- 1988). Rastrococcus invadens is usually scarce in
rus in Israel and the pest was controlled with the parts of India because it is controlled by natural
introduction of Clausenia purpurea (Ishii). enemies. The introduction of the encyrtid
Pseudococcus fragilis (Brain) was rst found in Gyranusoidea tebyi (Noyes) from India to West
California and rapidly became a serious pest of Africa, and its swift control of the mealybug
citrus to the point that it threatened the industry; there, is hailed as another biological control suc-
it also became a serious pest in Abkhazia of cess (Neuenschwander et al. 1994). Rastrococcus
USSR. iceryoides (Green) is causing serious damage to
Pseudococcus longispinus (Targioni Tozzetti) mango from India, and other fruit trees, and it is
is distributed worldwide. This mealybug is often also a pest of cotton. At present, it is distributed
11 Economic Importance 137
throughout India and eastwards to Thailand and bug, but it has caused signicant damage to orna-
Malaysia. It has also reached East Africa, where mental bougainvillea plants in Britain, ruining
there have been reports of damage in inland parts their aesthetic appearance and reducing their
of Tanzania and Malawi. Rastrococcus iceryoi- market value. Large mealybug populations cause
des is also known to cause serious damage to necrosis of the foliage, leaf loss, dieback and
Kapok trees in Tanganyika. Saccharicoccus sac- moulds grow on the excreted honeydew.
chari (Cockerell) is distributed wherever sugar- Mealybug is a pest, which can have a consider-
cane is grown, particularly Hawaii, Egypt, able negative economic impact on a wide range
Somalia, Costa Rica, etc. In southern Asia, it has of crops and ornamentals.
been rated as one of the most important mealy- There are some ground-inhabiting mealybug
bugs attacking sugarcane, which is the main species of undetermined economic importance
source of sugar and alcohol. It is also a possible belonging to the genus Rhizoecus and Geococcus.
vector of rice diseases in Cuba and India. Puto pilosellae (Sulc) is a pest of strawberries
Spilococcus mamillariae (Bouche) is a common (Kosztarab and Kozar 1988). These species are
pest of ornamental succulent plants. almost impossible to control. They are capable of
Trionymus radicicola (Morrison) caused causing serious economic damage to some crops,
severe damage to sugarcane in Cuba when areas namely alfalfa, strawberry, banana, pepper, cof-
of sugarcane dried out due to heavy population of fee, etc. Rhizoecus americanus (Hambleton) is
the mealybug on the roots. Trionymus townesi often a serious pest in Florida nurseries and
(Beardsley) is also known to attack rice and sor- recently it appeared in Italy, where it infests orna-
ghum. In the Philippines, infested upland rice mental plants. Paraputo leveri (Green) has been
crops have been reported to show depressed recorded as damaging roots and killing coffee
areas; plants in these areas were apparently plants in Papua New Guinea (Williams 1986a).
stunted and yellowish. Vryburgia rimariae Paraputo theaecola (Green) is found on tea roots,
(Tranfaglia) is a pest of economic importance in apparently in large numbers in North India. It is
greenhouses in Italy. also a severe pest on the roots of Taraktogenos
One of the most common groups of insects kurzii, a plant that produces a valuable oil.
attacking ornamental plants is mealybugs. There Paraputo banzigeri sp. lives on the roots causing
are about 275 species of mealybugs known to be the death of Dimopcarpus longan. Paraputo
present in the continental United States. leveri (Green) is already known from much of the
Mealybugs are prevalent pests in greenhouses tropical Pacic region and southern Asia. In
and interior plantscapes such as shopping malls, Papua New Guinea, it is found on the roots of
conservatories, hotels and ofce buildings. coffee, where it is protected under a layer of the
Mealybugs cost growers and retailers millions of fungus Diacanthodes philippinensis and eventu-
dollars per year in control and crop damage or ally kills the trees.
loss. Damage is caused by mealybugs feeding on Rhizoecus cocois (Williams) is a hypogeal
host tissues and injecting toxins or plant patho- species known from India, where it occurs on
gens into host plants. In addition, mealybugs coconuts, causing roots to dry up and young
secrete a waste product, honeydew, which is a plants to show loss of vigour. Rhizoecus dianthi
syrupy, sugary liquid that falls on the leaves, (Green) is a serious pest of African violets in
coating them with a shiny, sticky lm. Honeydew California (Snetsinger 1966) and a major pest of
serves as a medium for the growth of sooty mould greenhouse plants in Europe. Rhizoecus kondonis
fungus that reduces the plants photosynthetic (Kuwana) is one of the most widespread and eco-
abilities and ruins the plants appearance. Feeding nomically important subterranean mealybugs in
by mealybugs can cause premature leaf drop, die- California, where it is a pest of alfalfa, prune
back and may even kill plants if left unchecked. trees and strawberry plants and also caused
There is almost no information published on the severe damage to citrus in Japan. Rhizoecus
economic importance of bougainvillea mealy- amorphophalli (Betrem) was recorded from
138 M. Mani and C. Shivaraju
Trivandrum, Kerala (India), on the roots of ele- Some mealybug species may be manipulated
phant foot yam, Amorphophallus sp., ginger, as benecial insects in conservation biological
Dioscorea and rhizomes of Curcuma domestica control tactics. For example, the cupress mealy-
stored for seed purposes. Rhizoecus amorpho- bug, Planococcus vovae (Nasonov), which
phalli sucks the cell sap from the tubers, and occurs on cupress trees (Cupressus spp.) grown
severely infested deformed tubers of elephant in windbreaks, serves as an alternative host for
foot yam, taro, tannia nd no place in the market, natural enemies of mealybug pests in surround-
nor do they accept for cooking, causing economic ing citrus orchards and cocoa plantations (Cox
loss in India. 1989; Ho and Khoo 1997; Franco et al. 2004).
Rhizoecus americanus (Ferris) is a soft- Mealybugs have been also used as benecial
bodied, sucking insect that attacks the tips of insects in biological control of weeds. For exam-
roots. It is very common in Florida and other ple, Hypogeococcus pungens (Granara de
southern states. However, if shipped in plants, it Willink) was successfully introduced from
continues to thrive indoors and in greenhouses. Argentina into Queensland (Australia) for the
These creatures are dangerous to plants and are control of Harrisia cactus (Eriocereus martini)
often ignored as insignicant or misidentied as and related plants (Williams and Granara de
mycorrhiza. Willink 1992). In southern Asia, Trabutina ser-
Geococcus coffeae (Green) is found through- pentina (Green) is conned to Tamarix spp. in
out most of southern Asia, often killing plants in Pakistan and India. Heavy infestations of the
several counties, where it has been introduced. mealybug cause withering of the plant, and the
Heavy infestations of Geococcus johorensis mealybug has the potential for biological control
(Williams) in Malaysia cause the yellowing and of Tamarix wherever the plants have gained weed
early dieback of the leaves. Xenococcus acropy- status. The mealybug Hypogeococcus festerianus
gae (Williams) was found causing damage to the has been used to control Harrisia cactus, a major
roots of grapes in India. weed in central Queensland.
Several mealybug species are vectors of viral Mealybugs have also provided food for
diseases of various crops: banana, black pepper humans; the biblical manna, one of the food
(Bhat et al. 2003), cocoa (Dufour 1991), grapevine sources consumed by the Israelites during their
(Tsai et al. 2008), pineapple (Sether and Hu 2002), wandering in the wilderness of Sinai, is believed
rice (Abo and Sy 1998) and sugarcane (Lockhart to have been the honeydew excretion of the
et al. 1992). In such cases, mealybugs may be eco- manna mealybug Trabutina mannipara
nomic pests even at low densities. For example, (Hemprich and Ehrenberg) (Ben-Dov 2006;
several mealybug species are responsible for Miller and Kosztarab 1979).
GLRaV-3 transmission to grapevine, which has
been shown by the strong positive correlations
between mealybug numbers and infection levels in References
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Ecology
12
M. Mani and C. Shivaraju
areas, for they are normally found in the leaf or yuccae (Coquillett) or Puto decorosus McKenzie
stem axils. Infestations on fruits are usually usually just cling to the bark in any way possible.
enclosed in the calyx area. On the foliage, mealy- Mealybugs of the Phenacoccus type may also be
bugs such as Planococcus citri (Risso), found encysted in the centre of the trunk of the
Pseudococcus longispinus (Targioni-Tozzetti) perennial plant itself. Perhaps these pseudococ-
and P. obscures Essig are common. cids gain entrance into this area through cracks in
Stems and twigs of perennials are also good the bark, but they appear to be completely
areas for mealybug colonies. Normally, the enclosed by the plant tissue. Phenacoccus arte-
mealybugs are found wedged into the cracks and misiae Ehrhorn is an example of this type of
crevices of the bark, again showing a preference mealybug.
for tight areas. Stems and twigs of plants are not Very often, the plant parts, both above and
nearly as important as roots, crowns and leaf below the ground, are utilized by certain boring
axils, but mealybugs such as Spilococcus eriogoni insect larvae. Commonly, if these boring tunnels
(Ehrhorn), Phenacoccus delectus Ferris and P. are pulled open, heavy infestations of certain spe-
alleni McKenzie are occasionally found restricted cies of mealybugs will be discovered.
to the branches. Most often, branches are an Phenacoccus eremicus Ferris seems to depend
overow area from the heavily infested leaf axils. upon the cerambycid boring tunnels for its total
The habitat of the mealybugs may be divided existence during detrimental weather. When
into two parts: the part which is above the ground snow is on the ground or temperatures are consis-
(arboreal) and that which is below the ground tently above 100 F, these mealybugs are found
(hypogeal). Very heavy infestations often occur only in the cerambycid tunnels. Apparently, the
below the ground. Frequently, this area becomes tunnels are an ideal setting for this pseudococcid,
so heavily contaminated with mealybugs that for the empty cavities surrounding the beetle
they often overow onto the roots. It should be larva are quite often completely full of mealy-
noted that only very rarely does this overow of bugs in all stages of development. During more
soil-inhabiting mealybugs move to the plant parts favourable weather conditions, infestations out-
above the soil surface. Many species of mealy- side of the tunnels are quite common.
bugs are entirely restricted to the root region.
These mealybugs can be divided into two types:
those that inhabit the main, large roots, and those 12.2.2 Annual Flowering Plants
that restrict themselves to the small eshy roots.
Cracks and crevices are the common areas of Mealybugs infesting annual plants are unique in
infestation on the main roots, but on the eshy that their host is present for only a small part of
roots, infestations may occur in any area where the year. The location of these mealybugs during
there is a vacant space. Species commonly col- that part of the year when their host plant is
lected in the large root areas are Phenacoccus absent is still a question. This is still a mystery.
solani Ferris, P. artemisiae Ehrhorn, For example, a very heavy infestation of
Chorizococcus polyporus McKenzie, Puto paci- Phenacoccus eschscholtziae McKenzie was
cus McKenzie, Chnaurococcus trifolii (Forbes) found in early April on a large variety of annual
and many more. Species found in the second hosts. But no mealybugs were found on plant
group are usually representatives of the genus parts either above or below the ground after 2
Rhizoecus, with R. falcifer Kunckel d'Herculais, months. Probably the annual-infesting mealy-
R. gracilis McKenzie and R. kondonis Kuwana as bugs, the disappearance of the pseudococcid with
good examples. its host, are still a mystery. Infestations on annual
Infestations of moderate- to small-sized hosts are localized in the subterranean areas, and
mealybugs usually occur in cracks and crevices there are mealybugs that are commonly found on
on the bark, whereas large species such as Puto the foliage of such hosts.
12 Ecology 143
12.2.3 Grasses roots or the foliage of the conifer are the areas of
early development and the bark is merely a rest-
Mealybugs are found on grasses in three areas: ing place for later development where no feeding
the foliage, crowns and roots. Most of the mealy- occurs. In the case of Puto sandini Washburn,
bugs inhabit the leaf-blade sheath of the grass which infests Engelmann spruce, Picea engel-
plant. This type of pseudococcid is especially mannii, in Utah, at elevations of 10,00011,200
adapted to its habitat in that it is dorsoventrally ft, this mealybug goes through a very compli-
attened, allowing it to t comfortably in its cated 4-year life cycle involving annual multiple
sheath environment. Trionymus is the most com- migrations. As a demonstration of the complexity
mon genus of this type, with T. dolus Ferris, T. of the life cycle, here is what happens during the
smithii (Essig) and T. festucae (Kuwana) being rst year. The mature adult females produce the
good examples. Mealybugs are found on the leaf rst-instar nymphs under the bark chips on the
surfaces of the Gramineae. Again, Trionymus is a bole of the tree. These nymphs migrate from the
good example of this type of infestation. Some bole to the duff at the base of the tree in late
mealybugs are found in the crown regions of September where they remain under the snow
Gramineae, for example, Discococcus spp. Some until sometime in May. At this time, they migrate
mealybugs are found at the nodes of the grass back up the tree to the foliage where they feed
stems, for example, Antonina graminis (Maskell). until mid-July. At this time, they migrate back
Some of these mealybugs feed on the tender ter- down the bole and hide in the bark crevices. In
minal roots of the grass, where they are exposed September, many specimens go back to the foli-
to the surrounding soil with little or no waxy pro- age again where they feed intermittently until late
tection. Cryptoripersia, on the other hand, occurs September. They then move down to the ground,
in the same area on the roots, but is covered by a crawl into the duff where they remain until the
tough, felted sac. Another root area is inhabited following May. This type of migration seems fea-
by Discococcus spectabilis McKenzie. The sible for other conifer-infesting Puto because in
mature adult females of this mealybug form their some California species there are known records
ovisac at the base of the plant just a few millime- of infestations in the duff, on the bark and in the
tres below where the roots were rst produced foliage.
from the culm base. Areas other than the bark of conifers are also
infested. The foliage region is particularly well
inhabited. In most instances, the mealybugs are
12.2.4 Conifers found at the bases of the needles. As far as is
known, the total life cycle of these species is
The genus Puto contains several species which spent in the region of the foliage. Some species
inhabit various types of conifers. Most of them found in the foliage area are Crisicoccus pini
have been found only in the cracks in the bark or (Kuwana), Dysmicoccus pinicolus McKenzie, D.
under the rocks and fallen logs. This host posi- ryani (Coquillett), Ehrhornia cupressi (Ehrhorn),
tion is quite extraordinary because it would seem Puto cupressi (Coleman) and Spilcoccus implica-
virtually impossible for the mealybugs to gain tus Ferris. Root infestations in coniferous trees
any nourishment from such inanimate objects. are not common.
Some of these pseudococcids have been found on Fern-inhabiting mealybugs are not at all
the bark at the base of the trunks of redwood trees common. Rhizoecus pritchardi McKenzie is,
well over 150 ft tall. The bark in this area would however, a species which is often found on
undoubtedly be over ten times thicker than the the roots of maidenhair fern, and is perhaps
length of the mealybug stylets. more commonly associated with the roots of
Perhaps these late-instar nymphs were able to African violet. Several other mealybugs are
sustain themselves completely on nutrients gath- also found on, although not restricted to, vari-
ered in the early parts of their life history. The ous types of ferns in nurseries. Perhaps the
144 M. Mani and C. Shivaraju
most common species in this category is cially temperature and relative humidity (RH),
Pseudococcus longispinus (Targioni- are major ecological factors that have been found
Tozzetti). Several species of Pedronia are to severely affect mealybugs and their natural
restricted to ferns in Hawaii. enemies.
Occasionally, mealybugs are found in association Temperature is the driving force for mealybug
with nonbiological objects, especially during development. Many mealybugs overwinter as
winter. Misericoccus arenarius (Done and second-instar nymphs, although adult females,
Steinweden) is often found under the rocks or rst-instar nymphs and eggs also can full this
cracks in the rocks during the winter period. function (Miller 2005). For example,
Phenacoccus colemani Ehrhorn is commonly Phenacoccus azaleae Kuwana overwinters as a
found in the pits and crevices under the lava rocks. second-instar nymph within a wax cocoon (Xie
Heliococcus stachyos (Ehrhorn) is found under et al. 1999), Planococcus vovae as rst and sec-
the rocks deeply embedded in cracks in the rocks. ond instars (Francardi and Covassi 1992),
Wooden boards also serve as the habitat of some Pseudococcus viburni as rst instar in bark crev-
mealybugs. Chorizococcus rostellum (Hoke) is ices, and rarely as second or third instars
commonly found under the wooden boards. (Kosztarab 1996), and Pseudococcus maritimus
(Ehrhorn) as eggs and rst instar under the bark
(Geiger and Daane 2001). Ps. maritimus and Pl.
12.3 Host Plant Position cus overwinter primarily under the bark of the
trunk and cordon, with some of the population
Mealybugs show preference to certain positions found underground on the roots.
within the plant. Development of mealybug pop- The temperature has an impact on the devel-
ulation can be related to the plant growth and opment times and temperature thresholds for dif-
development. The mealybugs are relatively abun- ferent species. The number of generations of the
dant more on the fruits than on the other plant mealybug varies with the species, locality and
parts. This is true in several fruit crops. Mealybugs climatic factors. Most mealybug species are uni-
are found under loose bark and also on aerial or bivoltine, although some are reported to have
roots in the case of grapevine when fruit bunches as many as eight generations per annum in green-
are not available. Congregation of mealybugs is houses. There are ten generations of Planococcus
observed in the nodal region of the stem. They minor in a year, eight during FebruaryNovember
are seen usually on the lower surface of the and two during NovemberJanuary; M. hirsutus
leaves, more near the veins in the leaf. Mealybugs had ten generations per year in India. For exam-
are seen in large numbers on the fruits and are ple, Pseudococcus maritimus will have two gen-
rarely seen on the leaves and trunk in the case of erations in Californias interior valleys, whereas
custard apple. In ornamentals, the mealybugs are Planococcus cus can have seven generations in
distributed on the leaves, terminal shoots and the same region but is reported to have only three
ower buds. Some arboreal mealybugs extend generations per year in Italy. Similarly, Pl. citri in
their feeding on the parts just below the soil. Brazil has six generations per year in the south,
but up to 11 per year in the northeast where
grapes are produced throughout the year (two
12.4 Seasonal Development harvests per season). Other than Ps. maritimus
and H. bohemicus, there does not appear to be
Abiotic and biotic factors play a major role in the winter dormancy for the mealybugs. There is also
seasonal development of mealybugs besides the a variation in the seasonal feeding location and
phenology of the crop. Weather conditions, espe- movement on the plant among and within spe-
12 Ecology 145
cies, depending on factors such as regional tem- were registered at intervals of 2 h for 24 h on two
peratures and vineyard management practices, as separate days in July and September, respec-
described for Ps. maritimus and Pl. cus. The tively. In both the months, there was a high peak
mealybug population overwinters primarily of activity at sunset and a lower one at sunrise; in
under the bark of the trunk and cordon, with July, the peaks were separated by many hours of
some of the population found underground on the low catches, whereas in September the morning
roots, especially when tended by ants. There is no peak was much higher and the morning and eve-
diapause. On warm days, development may occur ning peaks were somewhat closer together than
during the winter months, with completion of the in July, owing to the shorter day length.
rst generation almost entirely under the bark. In tropical countries, the mealybugs occur on
From spring to summer, the Pl. cus population the plants throughout the year. Development of
follows the movement of plant resources from mealybug population can be related to the plant
roots to shoots to leaves. Four to ve generations growth and development. The mealybugs will be
are completed and population density can relatively abundant more on the fruits than on the
increase rapidly, although high summer tempera- other plant parts. The population is low in winter
tures, in excess of 40 C, may slow the growth of and rainy seasons and higher in summer months.
the population and increase mortality. As berries There was a highly signicant positive correla-
ripen and sugars develop, mealybugs move into tion of maximum and minimum temperature and
the berry clusters, rst attacking those near the a highly signicant negative correlation of morn-
vine cordon. The rapid population increase in ing and evening relative humidity and a nonsig-
summer is followed by an equally rapid decline nicant negative correlation of rainfall with
after harvest, resulting from biological controls mealybug population in vineyards in South India.
and abiotic mortality associated with high tem- They manage to survive under loose bark, feed-
peratures and vine senescence. ing at bases of spurs and callus tissue at the site of
The optimal temperature for populations of girdles in the off-season. In the absence of rains,
the cassava mealybug is between 20 and 30 there is a sudden spurt in the mealybug popula-
C. The cassava mealybug has poor survivability tion in dry seasons. Females of the mealybug
during rainy season because it is washed off the Glycycnyza turangicola are enclosed in a capsule
plant and drowns. The preferred temperature formed by hardened honeydew. This specializa-
range for the vineyard mealybug Planococcus tion is regarded as an adaptation to desert condi-
vitis (Nied.) was 1634 C, and this preference tions where humidity is extremely low in the
was unaffected by relative humidity. Linear Amudarya plain.
velocity increased with rises in the temperature. Heavy sporadic rains and cool temperatures of
Individuals that had been preconditioned at 95 % less than 20 C result in the temporary reduction
RH showed no consistent preference when pro- in the mealybug population. The pest population
vided with a choice of relative humidities, but build-up coincides with a high temperature of
those that had been desiccated avoided low 3040 C, low humidity (less than 40 %) and
humidities. The degree of humidity is probably fruit development.
perceived over the whole body. The mealybug is There is also a variation in the seasonal feed-
photonegative, and the reduced compound eyes ing location and movement on the vine among
are probably the photoreceptors. A rough sub- species and within species depending on regional
stratum was generally preferred to a smooth one. temperatures and vineyard management
Temperature appeared to be the main factor practices. There is no diapause, and slow devel-
determining the behaviour of the adults, and ori- opment may occur during the winter months
entation in relation to temperature, light, humid- under South Indian conditions. M. hirsutus was
ity and contact is mainly achieved by klinokinesis, active during winter also, without hibernation but
klinotaxis and orthokinesis. Catches of the male was most active during MarchOctober on roselle
mealybug, Pseudococcus calceolariae (Mask.), around West Bengal. In North India, the mealy-
146 M. Mani and C. Shivaraju
bug M. hirsutus on mesta overwintered in over- The ensete root mealybug Cataenococcus
lapping stages in the capsules of mesta beneath ensete Williams and Matile-Ferrero is a major
the persistent calyx and epicalyx of the old and pest in the ensete-growing regions of southern
dried plants and also in the soil during December Ethiopia. The ensete root mealybug, was
and January, when food plants were not avail- observed between 1054 and 2977 meter above
able. Mealybugs developed from a fraction of sea level (masl). Its infestation was severe only
third-generation eggs that did not enter winter between 1400 and 2200 masl. The highest infes-
diapause. tation (53.6 %) was recorded between 1600 and
Water-stressed plants have been reported to 1800 masl (Addis et al. 2010).
favour the increase in the mealybug population.
High nitrogen application enhances the results in
increase in the population of mealybugs. 12.4.2 Dispersal
In Java, Ferrisia virgata appears in large num-
bers during dry seasons of the year. It is more Adult male mealybugs are winged. The rst-
abundant from February to May in Philippines. instar nymphs (crawlers) have been found to
In Saudi Arabia, three generations were observed possess numerous characteristics that have been
in a year, early June, early July and August, and considered adaptations for dispersal behaviour,
the population increased with each generation. It including long legs and antennae (Gullan and
overwintered between January and early June as Kosztarab 1997). Adult males and newly
an adult female in the cracks and crevices of emerged rst-instar nymphs of most mealybug
trunks and branches and also on the fallen leaves. species display dispersal actively. Most mealy-
The female mealybug also migrated to the soil in bugs remain relatively stationary throughout
winter. The peak activity of F. virgata was their life (Miller 2005). However, some species
observed during JuneSeptember in West Bengal. move to different areas of the host for overwin-
On a number of host plants, it was most active tering, feeding, mating, ovipositing and moult-
during AugustNovember and MarchApril but ing (Franco 1994; McKenzie 1967; Miller 2005).
was very much reduced during December The occurrence of seasonal movements within
January (Mani and Krishnamoorthy 1993). the host has been reported for various mealybug
Several environmental factors inuence the pop- species, especially those associated with woody
ulation of F. virgata. Numerous records refer to plants such as Pl. citri, Ps. calceolariae, Ps.
heavy attack by F. virgata after prolonged viburni and Ps. longispinus in citrus (Franco
drought. The mealybug is favoured by dry 1994; Nestel et al. 1995); Pl. cus and Ps. mari-
weather in Java. The most important factor was timus in grapevine (Geiger and Daane 2001;
atmospheric humidity which exercises an indi- Godfrey et al. 2003); Pl. vovae in Juniperus spp.
rect effect through its inuence on parasitic fungi. (Francardi and Covassi 1992) and Ph. azaleae in
Temperature did not appear to have much effect bunge prickly ash (Xie et al. 1999). Franco
on F. virgata in Java. But a signicant positive (1994) suggested that immature feeding stages
correlation was found between the population of mealybugs on citrus tend to search for and to
density and daily maximum and minimum tem- settle at the major sinks, for example, growing
peratures in Saudi Arabia. Wind inuences the fruits, of the host plant in each phenological
dispersal and establishment of F. virgata in addi- period. We believe this hypothesis may also
tion by walking of the mealybugs. Windy areas explain the migratory movements of other
are highly susceptible to attack which is more mealybug species within various hosts. For
severe on hill tops than in valleys. F. virgata example, Pl. kraunhiae Kuwana was reported to
attacks the weak plants easily and spreads quickly feed on bacterium galls because they are sinks
on the younger plants exposed to sun and pro- for assimilates and have more nutritious phloem
tected from wind. It appears to prefer below sap and parenchyma than do normal plant tissues
5,000 ft altitude. (Yamazaki and Sugiura 2005). Other nymphal
12 Ecology 147
stages and adult female mealybugs are wingless, wild animals may aid the passive dispersal of
and some even legless; hence, they are not highly mealybugs (Kosztarab and Kozr 1988).
vagile, often xed to plants by their mouthparts Transport of nursery plants (ornamentals and
and always have a restricted distribution. fruit plants) infested with mealybugs aids the
However, the cassava mealybug (Phenacoccus spread of mealybugs from one location to
manihoti), rst reported in 1973 from the Congo another location.
areas of Kinshasa and Brazzaville, had become Among arthropods, ants have also been
established throughout the whole cassava belt of reported to disperse many mealybug species
Africa as far south as Mozambique by 1986 on (Gullan and Kosztarab 1997; Malsch et al. 2001;
the single plant genus Manihot (Herren and Ranjan 2006). Ants transported mealybugs from
Neuenschwander 1991). Mealybugs, particu- plant to plant between and within elds, thus
larly crawlers, having legs move limited dis- facilitating mealybug dispersal; adult females
tances within the plant and also between the and immatures of some mealybug species, asso-
plants (Kosztarab and Kozr 1988). Nevertheless, ciated with the ant genus Acropyga Roger, are
if conditions are favourable, crawlers usually carried by queens in their mandibles when found-
settle on the natal host plant, often close to their ing new colonies (Williams 1998). Associations
mother, which leads to an aggregative distribu- among invasive species of ants and mealybugs
tion (Gullan and Kosztarab 1997; Nestel et al. can be important in their success in new locations
1995). Similarly to most scale insects, crawlers (Helms and Vinson 2002). It is also observed that
are the mealybugs main dispersal agents, even mealybugs and other scale insects cling to locusts
though the mortality is very high (Gullan and during swarming. Misra (1920) reported another
Kosztarab 1997). First-instar mealybugs are eas- phoretic method of eggs and nymphs of
ily transported by the wind (Gullan and Maconellicoccus hirsutus (Green) being trans-
Kosztarab 1997). However, Williams and ported by nymphs and adult females of another
Granara de Willink (1992) reported that mealy- mealybug Ferrisia virgata (Cockerell).
bugs were believed to be distributed by air cur- All these methods are responsible for the local
rents over only short distances. In addition, and short-distance dispersal. Long-range disper-
dispersal strategies may be more passive, and sal of mealybugs is usually accomplished by the
crawlers of several species have been found to transport of infested plant material. Many species
exhibit behaviours that increase the chances of of mealybugs have been widely distributed by
wind dispersal and to use wind dispersal to commercial trafc, mostly carried on imported
migrate for several kilometres (Washburn and plant material (Williams and Granara de Willink
Washburn 1984). In India, Paracoccus margin- 1992). Papaya mealybug Paracoccus marginatus
atus spread very fast by wind dispersal of crawl- had spread very fast through the transport of
ers from the state Tamil Nadu to other states. fruits infested with mealybugs from Tamil Nadu
Crawlers from several species have also been to other states in India. Plants and their associ-
found to be able to survive without food for ated insects must have been carried to new areas
extended periods, which should again enhance by people for many centuries and people still
their dispersal success (Gullan and Kosztarab carry infested plant material, as shown by the
1997). Male and female nymphs do not differ in numerous quarantine interception records.
their dispersal behaviour or in their dispersal Because of their cryptic habits and small size,
success when dispersal is via crawler locomo- mealybugs are difcult to detect at borders dur-
tion. All the mealybug instars can be transported ing quarantine inspections, especially if their
on infested leaves blown by the wind. In addi- population density on plants is low (Gullan and
tion, water, bed-soil, humans and domestic and Martin 2003).
148 M. Mani and C. Shivaraju
Mealybugs are found attacked by various natural Syrphidae, Cecidomyiidae, Chamaeyiidae and
enemies in nature. The outbreak of mealybugs Drosophilidae (Diptera) are known to feed on
was observed in many instances with the applica- the mealybugs besides the spiders, mantids,
tion of broad-spectrum insecticides, which might ground beetles, assassin bugs, predatory stink
have disturbed the activity of natural enemies bugs, minute pirate bugs and predatory thrips.
particularly parasitoids and predators. This They are polyphagus feeding on a variety of
clearly indicates the importance of natural ene- mealybugs. Naturally occurring predators are
mies in the regulatory role of the mealybug popu- capable of suppressing the mealybugs on several
lation. In fact, there is a very rich natural complex occasions.
on arboreal mealybugs, but there is a poor natural
enemy complex, particularly natural predators or
parasites on root mealybugs. Withdrawal of 13.1.1 Coleoptera
insecticides results in the reappearance of natural
enemies, thereby regulating the mealybug popu- 13.1.1.1 Coccinellidae
lation. The natural enemies of the pests can be Both adults and larvae feed voraciously on all
divided into three categories depending on how stages of the mealybugs including the egg masses.
they feed on the pests. They are predators, para- The larvae of many predatory coccinellids are
sitoids or pathogens. covered with white waxy laments very similar
to the mealybugs. The adults are brightly
coloured. The eggs are oval shaped, yellow and
13.1 Predators very small. The larvae are voracious feeders
though the adults are also known to feed on the
Insects belonging to Coccinellidae (Coleoptera), mealybugs. Development from egg to adult bee-
Chrysopidae and Hemerobiidae (Neuroptera), tle takes 2530 days at 25 C. The species belong-
Lycaenidae and Noctuidae (Lepidoptera) and ing to genera Cryptolaemus, Brumus,
Aspidimerus, Stictobura, Orcus, Diomus, Nephus,
Sidis, Parasidis, Pseudoscymnus, Hyperaspis,
A.N. Shylesha (*)
ICAR-National Bureau of Agriculturally Important Scymnus, Sasajiscymnus, Exochomus,
Insect Resources, Bangalore 560 024, India Brumoides, Cleophora, Harmonia etc. are some
e-mail: anshylesha@gmail.com of the important predators of mealybugs. Among
M. Mani the coccinellids, Cryptolaemus montrouzieri
Indian Institute of Horticultural Research, Mulsant was extensively used to control a variety
Bangalore 560089, India
of mealybugs throughout the world.
e-mail: mmani1949@yahoo.co.in
Hyperaspis trilineata Mulsant can, during her lifetime, lay from 300 to 400
Hyperaspis trilineata Mulsant is a principal pred- eggs. The larvae are covered with white waxy
ator of Saccharicoccus sacchari (Cockerell). It is laments, very similar to that of mealybugs. The
reported to have a peculiar type of egg that is at beetle larvae are ufer and can run faster. The
rst at and resembles a whitey larva. They are larvae are often little in the crop because they are
laid singly and are hatched in 810 days. The very small and often found in the egg mass of the
young larvae feed for a time on mealybug crawl- mealybug. The eggs of mealybugs are oval
ers before developing their cottony covering. shaped, yellow and very small, but in practice not
About 3 weeks are required for larval develop- visible. Both adult beetles and larvae eat mealy-
ment followed by pupal development and adult bugs. When a mealybug is eaten, the dead
emergence. remains can be seen on trees as white uffy mat-
ter. The larvae mainly feed on eggs and young
Nephus includens Kirsch mealybugs. They can consume up to about 100
Nephus includens Kirsch is a predator of the cit- eggs per day or about 50 young mealybugs.
rus mealybug. Adult beetles are dark; they have
four orange/yellow spots on their backs. They are Scymnus coccivora Ayyar
about 2 mm in size. Its eggs are laid in the egg Scymnus coccivora Ayyar is known to feed on
mass laid by the mealybug. The female beetle several species of the mealybugs. Adults are light
13 Natural Enemies of Mealybugs 151
brown in colour measuring 1.7 1.3 mm in size. 1.2 and 5.2 days, respectively, and the total life
The pale yellow eggs are deposited singly in the cycle is completed in 23 days. The sex ratio is
colony of mealybugs. The grub has instars. Long 1:1. A single grub S. coccivora was known to
waxy strands develop on the later stage of the consume 308 eggs or 62 nymphs or 6.55 adult
grub. The pupa is oval and light brown in colour mealybugs (Mani and Thontadaraya 1987).
fringed with short brown hairs. The egg, grub,
prepupal and pupal stages occupy 4.1, 4.2, 9.5,
Mallada boninensis veins are forked. The larvae of the brown lace-
A single larva of M. boninensis was able to prey wings belonging to genus Hemerobius, and
about 345, 490 and 560 nymphs of Ferrisia vir- Sympherobius prefer to prey the early stage of the
gata, Planococcus lilacinus and P. citri, respec- mealybugs, though they are known to feed all the
tively (Mani and Krishnamoorthy 1990). three nymphal instars of the female mealybugs.
The rst-instar larvae of Sympherobius fallax
Chrysoperla carnea consume the second stage of the long-tailed
The larvae of the Ch. carnea grubs were found as mealybug P. longispinus more than any other
active predators on the mealybugs, and the preda- stages and did not eat the fourth (adult) stage,
tory grub preyed on all the stages of the mealy- while the second-stage S. fallax preferred the
bug. A single larva was known to consume 378 third-stage mealybugs. The third-stage S. fallax
eggs or 730 nymphs, or 95 adult females of P. also preferred the third-stage mealybugs. In the
citri. choice experiment, the rst-stage larval predators
preferred the second-stage mealybugs signi-
13.1.2.2 Hemerobiidae cantly more than the other two stages, while the
Hemerobiidae is a family of the Neuropteran second- and third-stage predators preferred the
insects commonly known as brown lacewings. third-stage mealybugs signicantly more than the
These insects differ from the somewhat similar second and the fourth stages. Darkness had a
Chrysopidae (green lacewings) not only by the marked effect on the feeding efciency of all
usual colouring but also by the wing venation. stages of S. fallax. The number of mealybugs
Hemerobiids have numerous long veins that are eaten in the light was signicantly greater (Gillani
lacking in chrysopids. Some of the costal cross et al. 2009).
The cilia of both the fore- and hindwings are of slightly paler brown. In the forewing, the cell and
the same shade as that of the ground colour of the apex are darker, with a white spot similar to that
wings. The antenna, head, thorax and abdomen in the male but larger, beyond the apex of the cell;
are pale brown in colour, and the club of the in most specimens, it is extended diffusely out-
antennae is ochraceous at apex; beneath are the wards and downwards. The hindwing is similar
palpi and thorax brownish-grey and the abdomen to that of the male. The underside is as precise as
pale brown in colour. In female, the upperside is in the male.
Second instar caterpillar Third instar caterpillar Final instar caterpillar Adult buttery
The lycaenid predator Spalgis epeus was com- ing to Drosophilidae, Cecidomyiidae, Syrphidae
monly associated with the natural control of the etc. are known to attack the mealybugs. The
mealybugs Rastrococcus iceryoides, Planococcus dipteran larvae feed voraciously on the
lilacinus, Pl. citri, Ferrrisia virgata, Paracoccus mealybugs.
marginatus etc. The larvae of Spalgis epeus were
observed to predate on root mealybug colonies 13.1.4.1 Cecidomyiidae
especially those at the base of the stems In this family, the larvae of a large number of spe-
(Devasahayam et al. 2009). Although Spalgis cies are predaceous on the mealybugs. These
lemolea was a common natural enemy of insects are very tiny, usually only 23 mm in
Phenacoccus madeirensis infesting cassava in length. The adults, which are very tiny, fragile
Africa (Herren and Neuenschwander 1991), its midges, locate colonies of appropriate prey. The
potential utility as an effective biological control eggs are laid near the base of the mealybug host;
agent was thwarted by its erratic occurrence. At the larva tunnel underneath the host and feed on
2530 C and 4060 % RH, the mean duration of the eggs or developing coccid nymphs. As the
the egg, larval and pupal stage of Spalgis epeus small, maggot-like larvae are incapable of mov-
on Pa. marginatus is 3.5, 12.0 and 10.3, respec- ing to considerable distances, there usually has to
tively, and the mean duration from egg to adult be a fair population of the prey present, before
emergence was 26 days, and it takes 24 days on the adults will lay eggs. The life cycle is com-
Pl. citri to complete the life cycle (Dinesh et al. pleted in 25 days. The total number of eggs
2010). As for the predatory potential of S. epeus, deposited by the female averaged 36 during her
the total number of papaya mealybugs consumed very short lifespan, which averaged 2.3 days. The
during the larval stage was 4,115 eggs, 281 larvae of Dicrodiplosis manihoti Harris were
nymphs and 77 female adults. found to prey on the egg masses of the cassava
mealybug, Phenacoccus manihoti in the Congo
and Senegal. Kalodiplosis pseudococci Felt has
13.1.4 Diptera given signicant control of D. brevipes in con-
junction with two parasitoids. Triommata coc-
Several dipterans are found as predators in the cidivora Felt plays a supplementary role in
concealed mealybug niche. The insects belong- regulating the mealybug population.
154 A.N. Shylesha and M. Mani
Pirate bugs feed on the mealybugs Crab spiders feed on the mealybugs
larva; it is usually composed of host blood cells and full-grown parasitoid emerges through an exit hole
the pigment melanin. The capsule may kill the para- at the distal part of the cocoon, leaving the lid
sitoid and thus prevent successful parasitism behind. Full development of the parasitoid takes
(Blumberg 1997). Various levels of encapsulation place inside the mealybug. Adult parasitoids feed
have been shown to occur in different mealybug themselves by piercing the young instars of the
species, in response to parasitism by encyrtids mealybugs and sucking from their bodies. By
(Blumberg 1997; Blumberg and van Driesche 2001; doing so, they can extend their lifespan. This feed-
Chong and Oetting 2007; Giordanengo and Nenon ing behaviour kills the young mealybug-instars.
1990; Sagarra et al. 2000). Conversely, encyrtid Parasitized mealybugs turn into a yellow/orange
parasitoids may use superparasitism as a strategy to cocoon and become hard (like mummies). These
overcome the immune response of unsuitable hosts mummies are difcult to see, because of their
(Blumberg et al. 2001). Besides superparasitism, small size. In this period, a female can lay about 80
other factors also affect the frequency of parasitoid eggs, most of them in the rst weeks of her life.
encapsulation including: (a) host and parasitoid spe- Anagyrus is a large genus of the family
cies; (b) the hosts physiological age and condition; Encyrtidae that attacks the mealybugs. Some
(c) the host and parasitoid origins (or strains); (d) important species like Anagyrus aegyptiacus, A.
the rearing and/or ambient temperature; and (e) the dactylopii, A. kamali, A. pseudococci play the
host plant species and stress conditions (Blumberg major role in suppressing the mealybugs. Other
1997; Calatayud et al. 2002). encyrtids, namely Leptomastidea abnormis,
Noyes and Hayat (1994) recorded 49 encyrtid Leptomastix dactylopii, Acerophagus papayae,
species as parasitoids of mealybugs in India. The Apoanagyrus lopezii, Aenasius bambawalei and
family Encyrtidae dominates the parasitoid com- Aenasius advena Comp., are found to be very
plex of mealybugs. Anagyrus, Apoanagyrus, effective parasitoids of mealybugs.
Adolescentus, Aenasius, Leptomastix,
Leptomastidea, Blepyrus, Gyranusoidea, Anagyrus antoninae (Timberlake)
Praleurocerus, Mahencyrtus, Acerophagus, It is an internal gregarious parasite of Antonina
Coccidoxenoides, Epidinocarsis, Neodusmetia, graminis. It is oriental in origin but common in
Hambletonia, Pseudaphycus and Alamella are Hawaii. It is active in cooler and high-humid areas.
some of the important genera under encyrtidae The female mates soon after the emergence and
attacking the mealybugs. They are sexually dimor- starts laying eggs immediately. Attack is on the
phic and both males and females are different from gravid female mealybugs. The stalked eggs are
each other. The males are smaller than the females unattached and free in the body uids of the mealy-
and have hairy antennae. The females have a bright bug and are hatched in 34 days. The larval and
band across the abdomen. The encyrtids are known prepupal stages cover 810 days. The pupal stage
to attack nymphs and adults of mealybugs. Each takes about 68 days, and the total life cycle is
species tends to specialize in terms of the stage of completed in 18 days. Up to seven adult parasites
development of the host. Certain species like emerge per mealybug and the sex ratio is 1:1. It is
Blepyrus insularis, Coccidoxenoides perminutus, carried out very well under Florida conditions.
Acerophagus papayae prefer earlier stage that is
58-day-old nymphs (early Second instar) for par- Neodusmetia sangwani (Subba Rao)
asitization, whereas species like Anagyrus dacty- It is an internal gregarious parasitoid of Antonina
lopii, Leptomastix dactylopii etc. prefer graminis and is native to India. Adult females are
1520-day-old mealybugs (third instar and young brachypterous and males are winged. They live
adult female). They breed very well when they are only for 2 days. The female produces up to 55
exposed to the preferred stage. The duration of the progeny. The sex ratio is 1:7. Life cycle is com-
life cycle is about 3 weeks at 25 C. Mealybugs pleted in 1723 days. Normal dispersal is very
that are parasitized turn into small cocoons, a little slow since the females are wingless. It has done
darker in colour than live mealybugs. The young very well under Texas conditions.
13 Natural Enemies of Mealybugs 159
Anagyrus fusciventris (Girault) bright blue eyes. Males are black in colour. Both
It is a parasitoid of Pseudococcus longispinus. sexes are about 3 mm in size. Mealybugs that are
Females are grey-brown in colour and have
parasitized turn into small cocoons, a little darker rate of three to four per day. The eggs hatch in 44 h
in colour than the live mealybugs. It prefers larger and the lifecycle is completed in 18 days at 27 C.
instars for parasitization. The females lay one egg
per host; from each parasitized mealybug, one Leptomastix dactylopii Howard
adult wasp will emerge. The lower temperature It is widely used against Planococcus citri.
threshold for the parasitoid is 18 C. The parasit- Besides P. citri, it also breeds well on Pl. cus.
oid development from egg to adult takes about 3 It is a small yellow-/brown-coloured parasitic
weeks at a temperature of 25 C. Full development wasp with distinctively long dark antennae. It is
of the parasitoid takes place inside the mealybug. about 3 mm long. Males are smaller and darker
Adult parasitoids feed themselves by piercing the than females. The antennae of the females are
young instars of the mealybugs and sucking from bended; the antennae of the males are hairy. Eggs
their bodies. By doing so, they can extend their life are laid in the third instar and in the young adult
span to about 2 months. This feeding behaviour female mealybug. The females deposit one egg
kills the young mealybug instars. inside the mealybug body. A female lays about 100
eggs. After hatching, the young larva of the parasit-
Anagyrus pseudococci (Girault) oid eats the mealybug from inside out. The parasit-
Anagyrus pseudococci (Girault) is native of ized mealybugs turn into a yellow-brown cocoon
Mediterranean areas. It is known to attack Pl. citri and become hard (like mummies). The lower tem-
and Ps. citriculus. It attacks all the nymphal stages perature threshold for Leptomastix dactylopii is
and the adult females but prefers the third instar of the 20 C, but the optimal temperature is 26 C. At
mealybug. About 45 eggs are laid per female at the 25 C, this development takes about 1517 days.
Leptomastix epona The adult parasite emerges from a circular exit hole
It is a parasitoid of Pseudococcus viburni (Ps. af- at the proximal end of the cocoon, leaving a lid on
nis) and Spilococcus cactearum. Adult wasps are the mummy. It mainly parasitizes older instars of
brown-black with thin, long, black antennae. Their Ps. viburni and Spilococcus cactearum. It lays one
wings are mainly translucent with slight dark bands. egg per mealybug. Mealybugs are killed by the
Leptomastix epona is 3 mm in size. Mealybugs that growing larva approximately 10 days after parasit-
are parasitized turn into yellow cocoon like mum- ization. Lower temperature threshold for
mies, easily distinguishable from live mealybugs. Leptomastix epona is 15 C.
26 C. Mealybugs are killed by the larva of the is a tiny small wasp with yellowish body, trans-
parasitic wasp, growing inside the mealybug. parent wings and grey/bluish eyes with three
Leptomastidea can survive temperatures up to black triangular spots in the forehead. The male
40 C. parasitoid is much smaller than the female para-
sitoid. This parasitoid affects mainly the second
Acerophagus papayae stage after hatching from the egg. Each female is
It is a solitary endoparasitoid of papaya mealy- capable of laying 50 eggs in its lifetime of 35
bug Paracoccus marginatus A. It parasitizes the days. Normally, single egg is laid inside a mealy-
early-stage (II instar) nymphs of the mealybug. It bug; occasionally more than one egg is also laid.
Gyranusoidea tebygi periods recorded were 18, 17, 16 and 14 days for
It is a native parasitoid of Rastrococcus invadens the rst, second, third and fourth nymphal instars,
Williams in India. The introduction of respectively. Oviposition commenced within
Gyranusoidea tebygi Noyes into Togo and Benin 24 h of emergence and lasted effectively for 6
was capable of eliminating R. invadens. It repro- days, during which 95 % of its eggs were laid and
duced on rst, second and third instars and it 1012 large hosts were killed through host feed-
avoided hosts that were already parasitized. Host ing. Sex ratio is 1:3.
feeding was occasionally observed. Sex ratios of
the offspring were male biased in smaller hosts, Aenasius bambawalei
as opposed to being female biased in larger hosts. It is a solitary endoparasitoid of Phenacoccus
Females had longer developmental times than solenopsis Tinsley in India and Pakistan. Egg and
males, developed faster in larger mealybugs than larval stages of the parasitoid are not visible
in smaller ones and were always larger than the being an internal feeder, but swelling and poor
males emerging from the same host instar. Their movement of the parasitized mealybugs were
size increased with the instar of the host at ovipo- observed after 23 days of parasitization. The
sition. About 90 % of all ovipositions in second- parasitized mealybugs transformed into dark-
and third-instar nymphs resulted from an attack brown mummies within 47 days. The pupae of
with multiple stings, starting with a sting in the A. bambawalei Hayat were barrel shaped with
head of the host for the most part. dark-brown colour. Duration of the pupal period
ranged from 5 to 8 days. Adults emerged by cut-
Apoanagyrus lopezii ting a circular small hole on the mummies after
Apoanagyrus (Epidinocarsis) lopezi (De Santis) completion of the pupal period. The adults of
is a species of the parasitic wasp native to Central both the sexes are shiny black in colour. Males
America. It is used as a biological control agent were smaller than females. The maximum devel-
against the cassava mealybug Phenacoccus mani- opmental time was recorded for the second-instar
hoti Matile-Ferrero in Africa. The parasitoid is host nymph as compared to the third instar. The
found to parasitize and complete development in males developed faster than the females in all
all developmental instars of Ph. manihoti. host stages. The overall sex ratio was 1:2. The
However, the parasitoid mortality was high maximum number of female wasps developed at
(15 %) when the development took place in the third-instar nymph (59.6 %), and it was con-
rst nymphal instar of the host. Complete devel- cluded that the third-instar host nymph appeared
opment from egg to adult emergence was pro- to be the most suitable host stage for mass
longed in smaller hosts, and the developmental rearing.
164 A.N. Shylesha and M. Mani
1290 days, respectively. The total life cycle of Adult aphelinids may feed on honeydew
A. japonica sp. n. is completed in 25.5 days. exuded by their hosts or on secretions issuing
Adults are small and short-lived. Longevity of from the wound caused during oviposition. The
the adults ranges from 7 to 11 days. The males eggs of aphelinids are often stalked. A number of
have long, hirsute, moniliform antennae, while endoparasitic species have an apneustic caudate
the females have shorter and distinctly clavate primary larva. Those that are endoparasitoids
antennae. Mating and oviposition takes place (e.g. Coccophagus) have larvae with neither spir-
readily. The adults exhibit a very good searching acles nor a functioning tracheal system. Some
capacity. A maximum of 238.16 parasitoids was species pupate inside the living host within a
obtained when the third-instar nymphs of 15 days pupation chamber, which becomes lled with air.
old were offered to A. japonica sp. n. for parasit- There is some evidence that the air inside this
ization (Mani and Krishnamoorthy 1989) chamber is derived from the hosts tracheal sys-
tem as in the Encyrtidae. Parasitoids emerge by
Allotropa utilis Muesbeck cutting a hole through the integument of the host
It is an internal, solitary parasitoid of nymphs of mummy; but if the mealybug has a delicate cov-
Phenacoccus aceris (Signoret) and Ph. pergandi ering, they push their way out from beneath it.
Ckll, native of Nova Scotia. It is reported to have The adults of some such species lack functional
a single generation. It attacks the smaller nymphs mandibles. Overwintering is normally as a
from July to October. Eggs laid in the body uid mature larva or pupa. The Aphelinidae are very
of the mealybugs increase sixfold during incuba- unusual in that the males and females may have
tion. Overwintering is by immature larvae in the different ontogenies. The females of such species
parasitized mealybugs. Pupation occurs in the always develop as primary endoparasitoids of
spring. The adult emergence takes place in May mealybugs.
from the overwintering host nymphs.
Coccophagus gurneyi
13.2.1.3 Aphelinidae It is quite polyphagous and is native of Australia.
Along with Encyrtidae, this family provides It is a solitary internal parasitoid of all the
most of the biocontrol agents. Aphelinids are nymphal instars of Ps. fragilis, Ps. comstocki and
small, soft-bodied parasitic wasps, yellow or Ps. longispinus. Coccophagus gurneyi Compere
brown in colour and do not typically exceed has a complex developmental biology. The
1.5 mm in length. The larvae of the majority are female-producing eggs are laid free in the body
the primary parasitoids on mealybugs. They are uids of the mealybug, where they hatch in about
found throughout the world in virtually all habi- 4 days at 27 C. The larva develops in 10 days
tats and are extremely important as biological followed by a 2-day prepupal stage and an 11-day
control agents. With regard to their biology, pupal stage. The total duration goes up to 44
Aphelinidae more closely resemble Encyrtidae. days. The male-producing egg of the parasitoid is
Characters uniting the family Aphelinidae are not deposited in the developing larva of the female
apomorphic; that is, they are not uniquely parasitoid. It gave a good control of Ps. fragilis in
derived. The characters of Aphelinidae are com- South Africa and Chile.
plete notaular lines of the mesoscutum; trans-
verse or broad petiole (propodeum); long 13.2.1.4 Other Families
marginal; short stigma; and short or absent post- There are species belonging to the families
marginal wing veins; and third valvula distinctly Braconidae, Eucoilidae, Signiphoridae,
separated and articulated with third valvifer. Eulopidae and Pteromalidae that are known to
These character combinations might also serve to attack the mealybugs but are of minor importance
differentiate Aphelinidae from other families of (Table 13.2).
Chalcidoidea.
166 A.N. Shylesha and M. Mani
The nematode Steinernema carpocapsae Chong JH, Oetting RD (2007) Specicity of Anagyrus sp
nov nr. sinope and Leptomastix dactylopii for six
(Weiser) was found to cause high mortality in
mealybug species. Biocontrol 52:289308
short time periods in adult females of the mealy- Clancy DW (1944) Biology of Allotropa burrelli, a gre-
bug Dysmicoccus texensis (Tinsley) (Andalo garious parasite of Pseudococcus comctocki. J Agric
et al. 2004). The aqueous suspension of EPN Res 69:159167
Devasahayam S, Koya KMA (2000) Evaluation of ento-
(JPM3) was more efcient with 70 % control
mopathogenic fungi against root mealybug infesting
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(Alves et al. 2009). Heterorhabditis bacteriophora Association for Advancement of Entomology,
Poinar strain HC1 was known to cause 100 % Trivandrum, 58 Nov 2000, pp 3334
Devasahayam S, Koya KMA, Anandaraj M, Thomas T,
mortality in the inoculated coffee mealybug com-
Preethi N (2009) Distribution and bio-ecology of root
plex (Rodriguez et al. 1998). Dysmicoccus texen- mealybugs associated with black pepper (Piper
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Greenhouse results demonstrate that the aqueous Entomon 34:147154
Dinesh AS, Venkatesha MV, Ramakrishna S (2010)
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Semiochemicals in Mealybugs
14
N. Bakthavatsalam
A semiochemical (from Greek semeon meaning 1978). In Italy, the sex pheromone released by
signal) is a generic term used for a chemical females of Planococcus citri (Risso) was
substance or mixture that carries a message for extracted from unmated females (Rotundo and
purpose of communication. Semiochemical com- Tremblay 1976). The identication of sex phero-
munication can be divided into two broad classes: mones of several mealybug species has facilitated
communication between individuals of the same the development of monitoring techniques and
species (intraspecic) or communication between management tactics based on these compounds.
different species (interspecic). It is usually used However, experience shows that the efciency of
in the eld of chemical ecology to encompass tactics such as mass trapping, mating disruption,
pheromones, allomones, kairomones, attractants, and lure and kill may be constrained by a lack of
and repellents. Many insects, including parasitic knowledge of basic features of the life history
insects, use semiochemicals, which are natural and mating behavior of male insects and the
chemicals released by an organism that affect the mechanisms involved in their interactions with
behaviors of other individuals. Pheromones are pheromone sources. A comprehensive account of
intraspecic signals that aid in nding mates, pheromones of coccoids was provided by
food and habitat resources, warning of enemies, Dunkelblum (1999) in the book Pheromones of
and avoiding competition. Interspecic signals Non-lepidoteran Insects.
known as allomones and kairomones have similar
functions.
The existence of female sex pheromone in a 14.1 Mealybug Pheromone
coccid, Matsucoccus resinosae Bean and Characteristics
Godwin, was rst demonstrated by Doane (1966),
followed by other coccid Aonidiella aurantii These pheromones all share a number of desir-
(Makell) (Tashiro and Chambers 1967). able characteristics and also some undesirable
Identication and synthesis of sex pheromone characteristics for use in pheromone trapping.
in A. aurantii became a turning point in the pher-
omone research for the coccoids (Roelofs et al. Sex pheromones are very powerful attractants
for male mealybugs. The males hide and live
in protected areas and are not harmed by the
N. Bakthavatsalam (*) insecticides. Using pheromones even small
National Bureau of Agriculturally Important Insects, populations can be detected at earlier stages of
Bangalore 560024, India occurrence.
e-mail: nbakthavatsalam@yahoo.com
The standard rubber septum-type pheromone pressure is high. When used in management
lures remain attractive for 23 months under systems, they are often combined with neonic-
eld conditions, minimizing the number of otinoids, insect growth regulators, or other
lure changes required throughout one season. biopesticides.
The lures require only a tiny dose of phero- Pheromone traps were used to determine, for
mone (0.025 mg or less), which will help to example, mealybug species composition,
keep lure manufacturing costs down. relative seasonal abundance, and density. In
The chemical structure of each pheromone is the vineyards infested with mixed mealybugs,
different; therefore, each pheromone speci- using traps activated with pheromones, it was
cally attracts only its own species. Sometimes found that Planococcus cus was more
in the orchards with more than one mealybug abundant than Planococcus citri in Italy (Ortu
species, pheromones could be combined to et al. 2006).
lure or to attract several species simultane- The ability of the pheromone control pro-
ously (Waterworth et al. 2011). grams to target a specic pest and not harm
Multi-season plastic Delta traps are sugested pollinator bees and benecial insects pays big
for monitoring and mass trapping most inva- dividends.
sive insect pests. This can also be an advantage and a disadvan-
Most pesticides require direct contact with the tage that mealybugs have been caught in traps
mealybug; hence, it will not be effective located over one quarter mile from the nearest
against those under the bark. Many of the known infestation.
softer insecticides are not able to penetrate The sensitivity of the multiplex polymerase
the waxy exterior; however, pheromones are chain reaction (PCR) to identify adult males
volatile molecules dispersed through the air collected in pheromone traps was shown, with
and sensed by the insect without requiring reliable identication of Pl. cus aged for 6
penetration-reach target spot. days in pheromone traps (Daane et al. 2011)
A possible alternative to visual sampling, which
may both decrease monitoring time and increase
the sensitivity of mealybug detection meth- 14.2 Techniques for Isolation
ods, is the use of pheromone-baited traps. of Pheromones
Specically, pheromone traps were more sensi-
tive than visual methods for detecting mealybug The pheromones of moths, due to their larger
infestations. The number of insects caught on size, were easily identied using the electrophys-
traps was correlated with mealybug abundance iological and analytical techniques; however, it is
in the eld indicating that pheromone traps difcult to identify the pheromones from mealy-
can be used in place of laborious, and annual bugs mainly due to their smaller size and avail-
sampling to monitor the mealybug populations. ability in lesser quantity of pheromones. Large
A second benet of the pheromone use over numbers of virgin females of mealybugs were
conventional broad-spectrum pesticides was required for the extraction of the female sex
that the ecological balance and natural preda- pheromone (Rotundo and Tremblay 1976). The
tor populations were preserved. Secondary large-scale production of mealybugs is done on
pest populations often surge later in the season potato sprouts or ripe pumpkins in several coun-
when broad-spectrum pesticides are used tries (Smith and Armitage 1920; Joshi et al. 2010;
because the pesticides kill natural predators Ahmad and Ghani 1970; Chacko et al. 1978).
of the primary pest, but with the use of phero- Behavioral attributes such as age of the calling
mones pest resurgence was contained. females and inuence of photo/scoto phases and
Vine mealybug pheromones are often inte- host plants complicate the process of pheromone
grated into pest management systems, particu- identication in mealybugs. For the isolation
larly for the rst several years when pest and identication of coccoid sex pheromones,
14 Semiochemicals in Mealybugs 175
continuous mass rearing of virgin females is might be the translucent pores on the hind legs
required, along with the sufcient numbers of coxae (metathoracic legs) (Gullan and Kosztarab
males for bioassays. Sexual separation of males 1997). The source of pheromone gland was sus-
was done either through the mechanical separa- pected to be the abdominal glands for several
tion using a water spray/brush or through the species of mealybugs. The pygidial glands were
application of juvenoids, which selectively pre- identied as the pheromone-producing glands,
vent male maturation. Pheromone production in the secretions of which are released through the
virgin females of Pseudococcus calceolariae was anus via a fragile duct. The production of phero-
not appreciably affected by the juvenile hormone mone and the response of the males to the phero-
treatment (Rotundo 1978). Compounds such as monal glands vary with the rearing conditions.
dichlorvos, triprene, RO 103108, etc. have been Feral mealybugs produce and respond to com-
used for the collection of females. pound 1, whereas lab-reared adults reared on
As a rst stage in research on the nature of potato sprouts produce and respond to both the
the female sex pheromone of the citrus pest compounds (Zada et al. 2003). For the collection
Pseudococcus calceolariae, a method was of pheromone, entrapment method using adsor-
described for collecting and extracting the phero- bent compounds such as tenax, Porapak Q or
mone from the air above virgin females reared on cold trapping was followed. The pheromone
potato sprouts in the laboratory; virgin females gland extracted with glacial acetic acid or abso-
sexed by insect growth regulators (which pre- lute ethanol was the most attractive pheromone
vented the development of males) and main- formulation for Planococcus citri (Hwang and
tained with or without food produced pheromones Chu 1987b). The plants that release volatiles
for about 16 days. Out of three absorbent materi- (such as lemon) need to be avoided as laboratory
als tested, Poropak Q proved to be the most effec- host to keep away the possible interference of
tive in absorbing the pheromone, which was these plant volatiles in the pheromone collection.
eluted from it with ethyl ether and concentrated Potato tubers or potato sprouts are considered to
by distillation or evaporation of the solvent in be the best rearing media for the collection of
nitrogen and the active fraction was subjected to pheromone.
gasliquid chromatography. All pheromone sam-
ples were evaluated by bioassay on the Poropak
in an olfactometer into which males were intro- 14.4 Behavior of Male Mealybugs
duced, or on lter paper in a petri dish containing
males (Rotundo et al. 1979a, b). In the course of The male mealybugs are known to y about one-
rearing Pseudococcus comstocki (Kuw.) on ripe half mile. The adult males usually respond to the
pumpkins in a laboratory in Japan, it was found pheromone gland secretions, and this was con-
possible to separate the sexes by the simple expe- rmed through electrophysiological and behav-
dient of wrapping the pumpkins on which the ioral studies. The daily rhythms of pheromone
mealybug was being reared in a double-folded production and the responsiveness of males were
sheet of tissue paper. Only the male nymphs very important. The males of P. citri were inac-
crawled into the paper. Adult males crawled out tive in scotophase but were very active for 30 min
from the papers and began to y soon after a after the beginning of photophase. However, the
lamp was lit (Negishi et al. 1980a). activity ceased after exposure to light. Besides
the adult males, immature stages of mealybugs
also respond to the pheromones of the female.
14.3 Pheromone Glands Second-instar nymphs of mealybugs (Pl. citri,
Ps. cryptus, N. viridis, and Pl. cus) responded to
The site of production and release of sex phero- the conspecic female sex pheromone directing
mones in female mealybugs have not yet been the males themselves to a suitable pupation site
determined, although it has been suggested that it near conspecic non-sibling mature females,
176 N. Bakthavatsalam
thus preventing in breeding. Male nymphs are Males of Pl. citri, Pl. cus, and Ps. comstocki are
not attracted to heterospecic female phero- morning iers, whose ight begins just after sun-
mones; the repellency of heterospecic sex pher- rise (Moreno et al. 1972, 1984; Ortu and Delrio
omones to males directed to look for a pupation 1982; da Silva et al.2009a, b, c; Zada et al. 2008);
site to avoid close contact with heterospecic males of Maconellicoccus hirsutus and
females (Mendel et al. 2003, 2012). Pheromotypes Nipaecoccus viridis (Newstead) y mainly
were also known to occur in Planococcus cus around sunset (Francis et al. 2007); and males of
(Signoret) (Kol-Maimon et al. 2010). The males Ps. calceolariae y both in the early morning and
of Pseudococcus calceolariae on citrus were able late afternoon (Rotundo and Tremblay 1976).
to move against the wind in a zigzag progression Moreno et al. (1984) suggested that the daily
(a klinotactic response), even in the absence of cycle of Pl. citri male ight activity is determined
pheromones. In still air, they responded to the by the scotophase period and its onset in response
pheromone source (a wad of cotton wool bearing to exposure to light. In light of the ndings of
100 u l of an ether extract of females), from Moreno et al. (1984) and O. Bar-Shalom and
within a radius of about 3 cm. In the presence of Z. Mendel (unpublished data), an endogenous
wind at the optimal speed (0.5 m/s) carrying the circadian rhythm, imprinted by the photoperiod,
pheromone, they moved directly toward the pher- may also be involved. Recently, Mendel et al.
omone source, even from a distance of over 1 m (2008) and da Silva et al. (2009a, b, c) studied
(a tropotactic response) (Rotundo et al. 1980). seven mealybug species of the genera
The turntable olfactometer method appeared to Planococcus, Pseudococcus, and Nipaecoccus to
be the most efcient to study the attractiveness of estimate for how long and at which physiological
the female sex pheromone of Planococcus citri age the mealybug males are sexually active.
to adult males. Forty 618-day-old females pro- Adult males take 3040 h to achieve sexual
vided a sufcient pheromone source, and red maturity before being able to y or to mate. Most
color sticky card was the most attractive. During mature males live for 23 days, during which
the scotophase, there was little ight by males; mating opportunities may be continuously avail-
they were attracted to the pheromone with an able, but searching for a mate by ight is limited
obvious peak within 30 min after the beginning to 24 h per day. The nding that mealybug
of the photophase. Male activity ceased 2 h after males appear to y only after exposure to day-
exposure to light (Hwang and Chu 1987b). The light suggests that visual cues also may be
responses of males to the extracted fractions involved in male ight and mate location.
showed that virgin females of both species Findings of recent experiments aimed at studying
Pseudococcus calceolariae and Planococcus the effects of the color and design of sticky traps
citri had no circadian rhythm of sexual activity baited with sex pheromone on male captures sup-
but emitted the pheromone consistently and con- port this hypothesis (Franco et al. 2008a). The
tinuously throughout the day until mated lack of simple eyes in apterous males of poly-
(Rotundo and Tremblay 1980). morphic species of mealybugs, such as S. sac-
chari (A 1968), is also an indirect evidence
that vision might be involved in male mealybug
14.5 Male Flight and Mate ight and mate location. As in other neococcoid
Location families, mealybug males typically have a pair of
dorsal and ventral simple eyes plus a pair of
In light of the pattern observed among the few smaller lateral ocelli (A 1968; Gullan and
species whose sex pheromones were identied, Kosztarab 1997). There is a lack of clear knowl-
mate location by mealybug males seems to rely edge about the functional aspects of this bizarre
mainly on chemical cues, that is, adult females of visual system. Duelli (1985) suggested that the
biparental mealybug species utilize sex phero- eyes in scale insect males are positioned in a
mones to attract males (Dunkelblum 1999). horizontal ring around the head because the
14 Semiochemicals in Mealybugs 177
males body axis is maintained almost vertical bug Pseudococcus viburni (Millar et al. 2005b),
during ight. The main purpose of the eyes would grape mealybug Pseudococcus maritimus
be to monitor the presence of females in conjuc- (Ehrhorn) (Figadre et al. 2007), passionvine
tion with pheromones. mealybug Planococcus minor (Maskell) (Ho
Unlike neotenic adult females, male mealy- et al. 2007), Japanese mealybug Planococcus
bugs are active iers, do not feed, and they live kraunhiae (Kuwana) (Sugie et al. 2008), long-
only a few days. The response of males of differ- tailed mealybug Pseudococcus longispinus
ent ages to a synthetic pheromone and virgin (Millar et al. 2009), Madeira mealybug,
females was tested. In the petri dish bioassay, Phenacoccus madeirensis Green (Ho et al. 2009),
class I males (up to 10 h after eclosion) and less citrophilous mealybug Planococcus calceolariae
than 20 % of class II males (1029 h after eclo- (Maskell) (El-Sayed et al. 2010) and Dysmicoccus
sion) responded to the pheromone or virgin grassii Leonardi (de Alfonso et al. 2012).
females. On the other hand, most of class III All known mealybug pheromones are mono-
males (29 or more hours after eclosion) showed a terpenoid esters, mostly of simple acids.
clear response. After eclosion, most P. citri males However, most of them are irregular non-head-
need to complete a period of sexual maturation of to-tail monoterpenoids, with unusual connections
at least 30 h before they can respond to the sex of two isoprene units (Millar and Midland 2007).
pheromone and mate. Without mating, the maxi- The majority of naturally occurring isoprenoid
mal lifespan of males was approximately 5 days compounds that have been identied have
and 50 % of males lived only up to 4.4 days 14,head-to-tail linkages between isoprenoid
(25.0 0.5 C). Most P. citri males have less than units, whereas most irregular terpenoids with
3 days to nd a receptive female and mate with non-head-to-tail linkages have been found in
her. However, since P. citri males only y within a members of the plant family Asteraceae (Rivera
period of approximately 4 h after sunrise, the total et al. 2001). Non-head-to-tail isoprenoid com-
effective time available for mate location by ight pounds are produced in three biosynthetic reac-
is only less than 12 h (da Silva et al. 2009a, b, c). tions, that is, cyclopropanation, branching, and
cyclobutanations (Thulasiram et al. 2008). Millar
and Midland (2007) suggested that terpenoid bio-
14.6 Identication/Isolation synthetic pathways in mealybugs are distinctly
of Pheromones different from the typical terpenoid pathways
found in other organisms, representing a variety
Using sophisticated analytical and bioassay of enzymes that can catalyze cyclizations and
instruments such as gas chromatographymass rearrangements. On this basis, and considering
spectral detector (GCMS), gas chromatography that mealybug endosymbionts are believed to be
electroantennogram detector (GCEAD), vibra- important to the nitrogen and sterol requirements
tional circular dichroism (VCD) spectroscopy, of their hosts and may play a role in physiologi-
and nuclear magnetic resonance (NMR) spec- cal processes such as resistance to microbial
troscopy, the sex pheromones of some economi- pathogens or detoxication of plant secondary
cally important species of mealybugs have been compounds, we tend to speculate that these
identied and synthesized. These include the enzymes may originate, at least in part, from
Comstock mealybug Pseudococcus comstocki mealybug endosymbionts. Thus, for example, a
(Negishi et al. 1980b), citrus mealybug variety of symbionts associated with bark beetles
Planococcus citri (Bierl-Leonhardt et al. 1981), are capable of producing compounds that are
vine mealybug Planococcus cus Signoret used as pheromones. Spectroscopically, phero-
(Hinkens et al. 2001), citriculus mealybug mones have been isolated and identied from
Pseudococcus cryptus Hempel (Arai et al. 2003), several species of mealybugs. A list of phero-
pink hibiscus mealybug Maconellicoccus hirsu- mones identied for several species is given in
tus (Green) (Zhang et al. 2004), obscure mealy- Table 14.1.
178
In Italy, the sex pheromone released by females A sex pheromone component of the Japanese
of Planococcus citri was extracted from unmated mealybug, Planococcus kraunhiae was isolated
females by ethanol, diethyl ether, or petroleum and identied. A crude extract of the pheromone
ether. Extracts in ethanol, diethyl ether, or petro- obtained by airborne collection was rst fraction-
leum ether placed on lter paper or hydrophilized ated with Florisil column chromatography. The
poly(methyl methacrylate) discs elicited high active fraction was further puried by HPLC, and
attraction and pairing responses in the males an active component was isolated by preparative
(Rotundo and Tremblay 1976, 1982). P. citri GC. The purified compound was determined
pheromone is a cyclobutane compound. to be 2-isopropylidene-5-methyl-4-hexen-1-yl
butyrate by GCMS and NMR analyses showing
the attraction activity to adult males of P.
14.6.2 Pseudococcus calceolariae kraunhiae in the eld (Sugie et al. 2008).
some eld tests it was as active as the latter (1R cis-3-isopropenyl-2-2-dimthyl cyclobutane
(Dunkelblum et al. 1987). The advantage of the methyl acetate) was found to attract males effec-
homolog is that its synthesis is easier and less tively (Dunkelblum et al. 1986). Alcohol analogue
expensive than that of the pheromone. Some (1R-cis)-3-isopropenyl-2-2-
pheromone analogs of the Comstock mealybug, dimethycyclobuanemethal) was an effective
Pseudococcus comstocki Kuwana, were also syn- attractant to P. citri, and homologue (1R-cis)-3-
thesized and tested in the eld (Uchida et al. isopropeny-2,2 dimethyl cyclobutane ethylacetate
1981; Bierl-Leonhardt et al. 1982). 2,6-dimethyl- at 2,000 g per dispenser was equal to 500 g
1,5-heptadien-3-yl acetate and three of its ana- pheromone (Dunkelblum et al. 1986). Analogue
logues of the sex pheromone of Pseudococcus of pheromone of P. citri, (+)-(1R)-cis-2-2-
comstocki (Kuw.), a pest of agricultural crops dimethyl-3-isopropenyl cyclobutano methanol
including apple and pear, were synthesized and acetate was synthesized using starting material cis-
evaluated for their attractiveness to males. All pinoic acid or cis-pinonic aldehyde, which were
four compounds were found to be the effective obtained from cheap -pinene and conversion of
attractants for the insect, but the synthetic sex the pinonic derivatives to pinononic derivatives
pheromone showed a two- to seven fold higher was achieved through Hundsdiecker reaction.
activity than the analogues (Uchida et al. 1981). Pinononyl aldehyde was used for synthesis of
pheromone through Wittig reaction (Dunkelblum
et al. 2002). Structural analogue of (+) cis-
14.7.2 Synthesis of Pheromones (1R)-(3)-isopropenyl-2-2-dimethyl cyclobutane
methyl acetate, the sex pheromone of P. citri, was
Through modications of acetoxy group, several synthesized and eld-tested in grapefruit orchards
pheromone analogues were synthesized for and the most active analogue was (+)-(cis-(1R)-
different species of mealybugs. A synthetic 2-(3-iso-propenyl-2-dimethyl cyclobutane ethyl
pheromone would provide a much more econom- acetate (Dunkelblum et al. 1987).
ical, convenient, and useful survey tool. Synthesis
of pheromone compounds of Pl. minor, Pl. citri, 14.7.2.2 Maconellicoccus hirsutus
and Ps. viburni was done successfully (Millar The sex pheromone of M. hirsutus, Maconelliol,
2008; Ho et al. 2007; Kukovinets et al. 2006; was synthesized in steps from Alpha pinene, and
Millar and Midland 2007). the key step was the dehydration of steps 57
through the intermediate 6 (Zhang et al. 2004).
14.7.2.1 Planococcus citri
The mealybug sex pheromones that have been 14.7.2.3 Pseudococcus viburni
identied generally are complex molecules, An improved diastereoselective synthesis of
which are relatively difcult to synthesize on a (1R*,2R*,3S*)-1-acetoxymethyl-2,3,4,4-
large scale. Nevertheless, because male mealy- tetramethylcyclopentane 1, the sex pheromone of
bugs are so exquisitely sensitive to the phero- Pseudococcus viburni, was described and the key
mone, with lures containing only a few step was diastereoselective catalytic hydrogena-
micrograms remaining active for at least several tion of the tetrasubstituted double bond in
months under eld conditions, widespread use of 2,3,4,4-tetramethyl-cyclopent-2-enone 4 to give
pheromone-baited traps for monitoring mealy- the thermodynamically less favored cis-2,3,4,4-
bugs is economically feasible. For example, 1 g tetramethyl-cyclopentanone 3a (Zou and Millar
of racemic pheromone is sufcient to prepare 2011). The pheromone of P. viburni was also syn-
50,000 lures or more @20 g per lure. thesized from pentalactone (Hajare et al. 2010).
In Pl. citri, (1R,3R)-3-isopropenyl-2,2- In the obscure mealybug Ps. viburni,
dimethylcyclobutanemethyl acetate (C12H20O2) 2,3,4,4-tetramethylcyclopentyl)methyl acetate
was identied, and a simple synthesis path was was identied as the sex pheromone. The active
developed in Israel, and the synthesized material compound has a number of isomers, and all were
184 N. Bakthavatsalam
made to conclusively verify the identity of the traps baited with virgin females. A mixture of
insect-produced compound. An efcient synthe- stereoisomers of the pheromone compound can
sis of the active compound, capable of being be used for eld trapping without adverse effects
scaled up to produce multigram quantities, was on trap catch (Unelius et al. 2011).
then developed. The pheromone was eld-tested
in California vineyards and nurseries, and by col- 14.7.2.5 Pseudococcus longispinus
laborators in South America and New Zealand. A single compound was unique to the headspace
The pheromone is extraordinarily active, with of the sexually mature female Ps. longispinus.
lures loaded with sub-milligram quantities The rst reported synthesis involves a polyphos-
remaining attractive to male mealybugs for sev- phoric acid-mediated cyclization of isobutyl
eral months. In South Africa, The sex pheromone 2-butenoate. The cyclopentenone was then con-
for P. viburni was recently identied and synthe- verted into the allylstannane after being reduced.
sized in South Africa. There was a positive and A short and efcient synthesis of the mealybug
signicant relationship between the fruit infesta- pheromone was developed from readily available
tion and number of P. viburni adult males caught iodoketone with an overall yield of 21 %. The
in pheromone-baited traps (r2 = 0.454, P < 0.001) pheromone has been shown to have extremely
in pome orchards. The action threshold level was high biological activity; in lures, just 25 g of the
estimated to be 2.5 male P. viburni caught per racemic pheromone can attract males for more
trap per fortnight at an economic threshold of 2 than 3 months (Bakonyi 2012). The synthesis
% fruit infestation. This monitoring method was of a recently identied and highly active sex
less labor-intensive, more accurate, and quicker pheromone of Ps. longispinus, was reported by
than the current visual sampling and monitoring Kurhade et al. (2013).
techniques (Mudavanhu et al. 2011).
14.7.2.6 Pseudococcus comstocki
14.7.2.4 Pseudococcus calceolariae The synthesis of the acetate of
Traps baited with 1001,000 g of racemic chry- 2,6-dimethylhepta-1,5-dien-3-olthe sex phero-
santhemyl 2-acetoxy-3-methylbutanoate cap- mone of the Comstock bughas been carried out
tured 420-fold more males than traps baited by condensing isobutenyl lithium with
with virgin females. In Chile, a single dose of 3,4-epoxy-2-methylbut-1-ene and acetylating
100 g was known to capture 1,171 males, the 2,6-dimethylhepta-1,5-dien-3-ol formed. The
whereas none were captured in control traps. An overall yield of pheromone was 46 % (Ishchenko
isomeric mixture of synthetic 3 proved to be et al. 1989). The synthesis of racemic versions of
highly attractive to male mealybugs in the eld in pheromones of Ps. comstocki was done through
New Zealand and in Chile. Male mealybugs were reductive lithiation of allyl phenyl thioesters
highly attracted to the racemic material and this followed by transmetallation, producing allylme-
will greatly facilitate the development of the tallics, which react selectively with carbonyl
pheromone for monitoring and control of this compounds at the most on least-substituted ter-
pest, because racemic 3 can be readily synthe- minus and the latter results in cis-olen
sized from commercially available intermediates (McCullough et al. 1991).
(El-Sayed et al. 2010). This activity of 1R,3R)-
[2,2-dimethyl-3-(2-methylprop-1-enyl)cyclopro-
pyl]methyl (R)-2-acetoxy-3-methylbutanoate in 14.8 Commercial Development
Pseudococcus calceolariae was further conrmed of Pheromones
by testing synthetic stereoisomers of the com-
pound as lures in traps for adult male mealybugs. Currently, of these seven pheromones, only the
Traps baited with 1,000 g of the pheromone vine mealybug Pl. cus pheromone is commer-
compound caught 36 times more males than cially available, but Millar is working to transfer
14 Semiochemicals in Mealybugs 185
the manufacturing technology to companies that Place the rubber septum containing the phero-
produce pheromone products. None of the phero- mone (lure) inside the trap on top of the sticky
mones are protected by patents; therefore, all are coating on the bottom panel.
freely available for the commercial development.
Companies also need to know that there is a sub- 14.9.1.2 Trap Placement
stantial market for these products, so growers Tie the trap to the plant at 23 feet above
should communicate their needs to company rep- ground level. Traps baited with virgin
resentatives to expedite the entry of mealybug Pseudococcus comstocki females, placed in
pheromone traps into the market place. Ps. comstocki-infested fruitless mulberry
Chemist Aijun Zhang in Beltsville, MD, trees about 9 ft above ground level, had
developed the pheromone of M. hirsutus, which caught an average of 225 Ps. comstocki
mimics the female mealybugs scent, according males compared with 6 ft by each of the
to a news release. South Carolina Scientic Inc., other traps (Moreno et al. 1972). Make sure
of Columbia, SC, will market the chemical. The leaves and shoots are not obstructing the
sex pheromone, placed inside sticky traps, effec- entrance into the trap. Do not hang it too low
tively monitors and traps male mealybug. By lur- or too high in the canopy.
ing males to traps, the pheromone would provide Place the trap at the center of the block for
a much more useful detection tool. Relatively surveying the largest area possible.
high concentrations of the pheromone repel
males away from the source, disrupting mating. 14.9.1.3 Labeling Traps
However, natural enemies of the mealybug are Label the trap with the block name and
not attracted to the scent, so biological control row number, where the trap was placed
would not be compromised (http://www.the- and the dates it was set out in the field and
grower.com/news/rm_to_market_pink_mealy- removed.
bug_ pheromone_117886474.html#sthash. Label the outer side of the trap with the fol-
2KIGfvcv.dpuf). Commercial lures of Ps. longis- lowing information: date of placement (DOP),
pinus and Ps. maritimus also became available vineyard and block name, row and vine num-
from Suterra LLC (Bend, OR) in 2010. ber, and lure (L) type. When you remove the
trap, write the date of removal (R). Use a per-
manent marker.
14.9 Traps
14.9.1.4 Trap Density
Pheromone traps can attract the mealybugs Place one trap per hectare or one per smaller
within one-quarter mile from the trap site. orchard.
especially in relation to trap design, trap color, Information on the level of male capture in spring
type of dispenser, pheromone dose, and bait lon- or early summer by application of pheromone
gevity and range (Francis et al. 2007; Franco traps is used to predict mealybug density or per-
et al. 2004a, 2008a, 2009; Millar et al. 2002; centage of fruit infestation and consequently to
Vitullo et al. 2007; Walton et al. 2004; Zada et al. assist in the decision making for the purpose of
2004, 2008). Nevertheless, the use of pheromone the citrus mealybug management (Franco et al.
traps as a monitoring tool for mealybug damage 2001). The analogous compound, 1R cis-3-
risk assessment depends on the existence of a isopropyl-2-2-diemthyl cyclobutyl methyl acetate
reasonable relationship between the number of was impregnated at 2,000 g in each dispenser
males captured in pheromone-baited traps and almost equaling 500 g of pheromone of Pl. citri
other mealybug infestation parameters, as to monitor the mealybugs (Franco et al. 2004b). In
recorded by other means, usually, visual sam- Mediterranean Basin, the best time for releasing
pling. A linear relationship was found to exist the natural enemies Cryptolaemus montrouzieri
between the vine mealybug, Pl. cus (Walton Muls. and Leptomastix dactylopii How. for the
et al. 2004), and the citrus mealybug, Pl. citri control of Pl. citri in citrus orchards was deduced
(Franco et al. 2001). However, this correlation from mealybug population monitoring by means
may be affected by different factors, including of traps containing the sex pheromone of P. citri
the weather, the activity of natural enemies, and (Panis 1981).
the phenological gap between male captures and
infestation level (Franco et al. 2001, 2008a). 14.10.1.3 Pseudococcus longispinus
and Pseudococcus viburni
14.10.1.1 Pseudococcus comstocki Positive and signicant relationship exists
The seasonal ight activity of Ps. comstocki in between pome fruit infestation and number of Ps.
California was monitored with sex pheromone vulbuni in pheromone traps baited with phero-
traps (Meyerdirk and Newell 1979). In the mone, and pheromone monitoring was done to
Moldavian area of the USSR, a synthetic material estimate the action threshold, which was esti-
of sex pheromone proved very successful in mated to be 2.5 male Ps. viburni (Mudavanhu
detecting not only the presence of Ps. comstocki et al. 2011). Lures containing 25 g per lure was
but also the information on its population density attractive to Ps. viburni and Ps. longispinus.
in mulberry ecosystem (Bichina et al. 1982). Racemic mixtures of S-lavandulyl senecioate and
Pheromone of Ps. comstocki containing (S)-lavandulyl isovalerate recorded good capture
2,6,-dimethyl-1,5,-heptadient at the rate of 200 (Zada et al. 2008). Operational parameters of
g per trap, placed at a height of 1.52 m, was able traps baited with the pheromones of three
to trap the adult males, and used for recording the mealybug species were optimized in nurseries
uctuation of daily and seasonal ight dynamics producing ornamental plants. All pheromone
(Smetnik and Rozinskaya 1988). doses (1320 g) attracted Ps. longispinus and
Ps. viburni males, with the lowest dose (1 g)
14.10.1.2 Planococcus citri attracting the fewest males for both species.
Monitoring population densities of Planococcus Lures containing 25-g doses of either phero-
citri in citrus ecosystem was based on male mone had effective eld lifetimes of at least 12
capture using traps baited with female sex phero- weeks. Pheromone traps were used to detect
mones. Pheromone of Pl. citri was used for the infestations of Ps. longispinus throughout the
early detection of the pest occurrence in citrus season and to track population cycles. When
elds (Ortu and Delrio 1982). In Israel, the sea- pheromone-baited traps for Ps. longispinus were
sonal population uctuations and trends of Pl. compared with manual sampling, trap counts of
citri were monitored (Hefetz and Tauber 1990; male mealybugs were signicantly correlated
Tauber et al. 1985) and traps were being used in with mealybugs counted on plants in the vicinity
conjunction with biological control methods. of the traps (Waterworth et al. 2011).
14 Semiochemicals in Mealybugs 189
synthesized in one step from two commercial ping, but not signicant enough to cause any
starting materials, so that it can be prepared in reduction in the infestation on fruits in
large quantities, sufcient for eld work, includ- Mediterranean countries (Franco et al. 2003).
ing mating disruption (Ujita and Saeki 2008; Similarly in Israel and Portugal, signicant
Walton et al. 2006). When the pheromone was reduction of male numbers can be achieved by
applied to the leaves as a sprayable microcapsule mass trapping with sticky plate traps (30 cm 30
formulation, crop damage was reduced from 9 to cm) baited with 200 g of pheromone used at a
11 % in control plots to 34 % in treated plots; rate of one per citrus tree, although fruit infesta-
however, the effective life of the formulation tion did not reduce signicantly. Therefore, as
presents a technical problem that needs to be the pheromone trapping system used cannot
solved. The efciency of the pheromone formu- reduce the number of attracted males effectively,
lation in the eld declined after 3 weeks, indicat- it is most likely that many of them originated
ing that more than four applications per season from outside the subplot. In fact, males are
were needed. The proximity of the mealybug attracted to the pheromone source from ranges up
sexes on emergence may also impair the success to at least 100 m (Branco et al. 2006; Franco et al.
of mating disruption (Walton et al. 2006). Mating 2004a). On the other hand, the higher level of
disruption in Planococcus kraunhiae through the mating observed in mass-trapping plots early in
use of pheromone, 2-isopropyliden-5-methyl-4- the spring, when the mealybug density is usually
hexenyl butyrate controlled the mealybug popu- very low, suggested that mass trapping led to a
lations in the eld in Japan (Teshiba et al. 2009). strong attraction of males from outside the sub-
There was a lesser density of Planococcus cus plots. In light of this nding, it was postulated
on leaves of vines treated with plastic dispensers that early in the season, when the male popula-
with 100 mg each of synthetic sex pheromone tion is usually low, the attraction of males to the
than the control; however, the difference was not edge of the orchard by using attractannihilate
signicant (Cocco et al. 2011). Due to its effec- tactics combined creates a male vacuum inside
tiveness in traps, developing the pheromone to the plot and, consequently, reduces mating and
control vine mealybug populations (Planococcus infestation (Franco et al. 2004a).
cus) using mating disruption was pursued
(http:// advancinggreenchemistry. org/ catch-all/ 14.11.2.2 Maconellicoccus hirsutus
mating-disruption-as-a-pest-management-tool- The quantum of pheromones produced through
in-californias-wine-industry/). synthesis was not sufcient to use for mass
trapping and mating disruption. However
recently, Chemist Aijun Zhang in Beltsville, MD,
14.11.2 Mass Trapping developed the pheromone, which mimics the
female mealybugs scent, and South Carolina
An articial lure might also enable the development Scientic Inc., of Columbia, SC, will market the
of mass trapping and mating disruption technology chemical. There is also a second potential control
for managing this pest, which would complement strategy (http://www.thegrower.com/news/rmto
the ongoing biological control eradication efforts. market pink mealybug pheromone_117886474.
In addition to their use for detection and monitoring html#sthash.2KIGfvcv.dpuf).
of insect populations, pheromones also have a
potential use in insect control, for example, by 14.11.2.3 Pseudococcus calceolariae
mating disruption or attract-and-kill technologies. In 1975, a mass-trapping experiment was carried
out against Pseudococcus calceolariae (Mask.),
14.11.2.1 Planococcus citri which was causing heavy damage on citrus near
Pheromones are yet to be exploited to a great Salerno, Italy; 79 traps baited with 1,538 virgin
extent for mass trapping of mealybugs. There females were placed in 25 orange trees over an
was a signicant reduction in the population of P. area of 1 ha in a single orchard, and they caught
citri when pheromone was used for mass trap- about 300,000 males of the species. Populations
14 Semiochemicals in Mealybugs 191
were sampled in 19751978 in order to assess the The sex pheromone of mealybugs may be
long-term effect of the traps on population used by their natural enemies as a kairomonal cue
dynamics. Up to 1976, a sevenfold reduction in in host or prey selection. Anagyrus pseudococci
captures of males and a tenfold decrease in popu- sp.n. is an effective parasitoid of vine mealybug
lations on fruits in this orchard were registered. Pl. cus and citrus mealybug Pl. citri. Anagyrus
Flight peaks of Ps. calceolariae occurred in mid- pseudococci in California vineyards was attracted
May, mid-July, and late September. It was not to the pheromone ((+)-lavandulyl senecioate
possible to determine from the experiments (LS)) of Pl. cus (Millar et al. 2002; Franco et al.
whether the decline in Ps. calceolariae was due 2008c) but Anagyrus pseudococci sp.n. was not
to either the earlier catches of males in the traps attracted to the pheromone ((+)-(1R,3R)-cis-2,2-
or sudden increases in populations of coccinellid dimethyl- 3-isopropenyl-cyclobutanemethanol
predator (Rotundo et al. 1979a, b). acetate (PcA, namely, planococcyl acetate) of P.
citri (Franco et al. 2008c, 2011; da Silva et al.
14.11.2.4 Planococcus cus 2009a, b); and this kairomonal response was an
The use of synthetic sex pheromones, such as innate behavior trait. An interesting aspect of this
those found in Suterras CheckMate products program is that a parasitoid of the vine mealybug
for mating disruption of vine mealybug, aims to (Anagyrus pseudococci) may be attracted to the
prevent adult males from mating. The use of mealybug pheromone as a host-nding cue,
degree-day models, pheromone traps, and eld resulting in greater levels of biological control.
observations are helpful for detecting the earliest There is a minimal risk of parasitoids being
colonies of mealybugs. By preventing mating caught if lures are deployed in triangular tent-
and subsequent egg laying, vine mealybug popu- shaped Delta traps. It was also found that the
lations can be dramatically reduced to below presence of Pl. cus sex pheromone signicantly
economically damaging levels. increases the parasitization rate of Pl. citri colo-
nies by Anagyrus pseudococci. (Franco et al.
2008b). In eld trials in Portugal, Italy, and
14.11.3 Kairomonal Response Israel, the rate of parasitism by A. pseudococci
was improved through the use of pheromone
The pheromone-lled air also acts as kairomones (Franco et al. 2001). In Sicilian orchards (Italy)
for several predators and parasitoids. The sex infested with Pl. cus, the number of captured A.
pheromone emitted by mealybug virgin females pseudococci females per trap was signicantly
provides reliable information on the location of a higher in LS (sex pheromone (S)-(+)-lavandulyl
potential host for mealybug parasitoids, because senecioate (LS)-baited traps resulting in the
of the sedentary nature of mealybugs. enhancement of the parasitoid performance
Furthermore, because of the typical clumped spa- (Mansour et al. 2010).
tial pattern of mealybugs, the sex pheromone will Pheromone-based mating disruption of vine
also be a convenient chemical cue by which the mealybug indicated that the treatment had no
parasitoid can efciently locate aggregates (colo- negative effect on the level of parasitization
nies) of hosts, which are expected to emit a stron- (Walton et al. 2006) with Pl. cus by A. pseudo-
ger pheromonal signal than that of single virgin cocci. The kairomonal response of Anagyrus
females (functional response). Thus, sex phero- pseudococci sp.n. could be explored in connec-
mones of mealybugs could serve as a novel and tion with biological control tactics, by enhancing
efcient tool to support the classical biological parasitization of Pl. citri as a component of inte-
control of invasive mealybug species, by identi- grated pest management strategies, by means of a
fying, in the region of origin of the target species, similar approach to that used against aphid pests
parasitoids that could be the potential candidates (Powell and Pickett 2003). Rotundo and
for use in the biological control program (Franco Tremblay (1975) reported that traps baited with
et al. 2008c). virgin females of Ps. calceolariae captured sig-
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Pseudococcidae) within the framework of integrated dimethylcyclobutane (sex pheromone of the citrus
control in Mediterranean citrus-growing [French]. Rev mealybug) from verboxide. Ser Khim 7:16031607
Zool Agric Pathol Veg 78(3):8896 Serrano MS, Lapointe SL, Meyerdirk DE (2001)
Passaro LC, Webster FX (2004) Synthesis of the female Attraction of males by virgin females of the mealybug.
sex pheromone of the citrus mealybug, Planococcus Maconellicoccus hirsutus (Hemiptera:
citri. J Agric Food Chem 52:28962899 Pseudococcidae) Environ Entomol 30(2):339345
Powell W, Pickett JA (2003) Manipulation of parasitoids Shaw PW, Walis DR (2011) Pheromone trap and crop
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Pest Manag Sci 59:149155 apple orchards. N Z Plant Protect 64:291
Rivera SB, Swedlund BD, King GJ, Bell RN, Hussey CE, Shaw P, Wallis D, Charles J (2012) Recent records of the
Shattuck-Eidens DM, Wrobel WM, Peiser GD, mealybug parasitoid Alamella mira Noyes
Poulter CD (2001) Chrysanthemyl diphosphate syn- (Hymenoptera: Aphelinidae) captured in citrophilus
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recombinant non-head-to-tail monoterpene synthase mone traps in Nelson. The Weta 42:3437
from Chrysanthemum cinerariaefolium. Proc Natl Skatteboel L, Stenstroem Y (1989) Facile synthesis of
Acad Sci U S A 98:43734378 racemic 3-acetoxy-2,6-dimethyl-1,5-heptadiene, the
Roelofs WL, Gieselmann MJ, Carde AM, Tashiro H, sex pheromone of the Comstock mealybug. Acta
Moreno DS, Henrick CA, Anderson RJ (1978) Chem Scand 43:9396
198 N. Bakthavatsalam
Smetnik AI, Rozinskaya EM (1988) A new method is Pseudococcidae) in nurseries producing ornamental
introduced. Zashchita Rateni Moskva 3:4243 plants. J Econ Entomol 104(2):555565
Smith HS, Armitage HM (1920) Biological control of Wolk JL, Goldschmidt Z, Dunkelblum E (1986) A short
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Bull 9:104158 tate, sex pheromone of the citrus mealybug
Sugie H, Teshiba M, Narai Y, Tsutsumi T, Sawamura N, Planococcus citri (Risso). Synthesis 4:347348
Tabata J, Hiradate S (2008) Identication of a sex Zada A, Dunkelblum E (2006) A convenient resolution of
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Zool 43(3):369375 tiomerically pure (R)-lavandulol. Tetrahedron-
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matsumotoi. Naturwissenschaften 99(7):567574 alcohols and of the two enantiomers of lavandulyl sene-
Tashiro H, Chambers DL (1967) Reproduction in the cioate. Tetrahedron-Asymmetry 15:23392343
California red scale, Aonidiella aurantii (Homoptera: Zada A, Dunkelblum E, Assael F, Harel M, Cojocaru M,
Diaspididae). I Discovery and extraction of a female sex Mendel Z, Pellizzari G (2001) Identication of a sec-
pheromone. Ann Entomol Soc Am 60(6):935940 ond sex pheromone component of the vine mealybug.
Tauber O, Sternlicht M, Hefetz A (1985) Preliminary Boll Zool Agrar Bach 33(275):281
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Ant Association
15
M. Mani and C. Shivaraju
The classic antaphid mutualistic relationship were also observed on grapevines in Europe in
has long been observed by naturalist and ento- association with Phenacoccus aceris (Signoret;
mologist alike, and several studies were con- Sforza 2008).
ducted on the actual/benets and factors involved
in these associations. This type of relationship
between ants and other insects is known to occur 15.1 Benets to Mealybugs
in a number of homopterous groups, especially in
the mealybugs. In the case of mealybugs, the Benets derived by mealybugs are more numer-
degree of dependence on the ants may vary from ous than might be expected.
strong and almost necessary associations to
weak, casual seasonal relationships. The associa-
tion of ants with the mealybugs resulted in the 15.2 Protection from Natural
hypothesis more the ants, more the mealybugs. Enemies
Ants are often associated with mealybugs as hon-
eydew consumers. Hemiptera-tending ants are Ants protect mealybugs from their natural ene-
mostly species of the subfamilies Myrmicinae, mies; this is a very important and long-realized
Dolichoderinae, and Formicinae (Degen and aspect of mealybug benet. Natural enemies are
Gersani 1989; Mittler and Douglas 2003). easily disturbed by movements of the ants. Ants
Samways et al. (1982) reported that 11 % of the are naturally hostile to any quick or obviously
123 ant species identied in citrus orchards in harmful movements around the honeydew
South Africa were associated with mealybugs. sources (Herzig 1938; Nixon 1951). The disrup-
Some mealybugs have an obligatory association tion of the activity of natural enemies by ants pro-
with ants: all Southeast Asian myrmecophilous vides a temporal refuge for mealybugs (Gutierrez
mealybugs have been collected only with ants of et al. 2008). Ants have long been known to aggra-
the genera Acropyga, Dolichoderus, or vate mealybug populations and other honeydew-
Polyrhachis, which attend the mealybugs either producing insect species by disrupting the natural
in subterranean nests or on aerial plant parts biological controls on these species.
(Gullan and Kosztarab 1997). Aboveground nests Ants deter the natural enemies of mealybugs.
There are numerous examples of ants deterring
M. Mani (*) C. Shivaraju the predators and parasites of mealybugs. For
Indian Institute of Horticultural Research, instance, ants also reduce parasitism of the cas-
Bangalore 560089, India sava mealybug, Phenacoccus manihoti
e-mail: mmani1949@yahoo.co.in
Matile-Ferrero (Cudjoe et al. 1993). A wide vari- attending the sucking pests. In Kenya, C. mon-
ety of natural enemies are known to prey on pine- trouzieri, released for the control of Planococcus
apple mealybugs. Ants protect mealybugs from kenyae (LePelley), was eliminated by ants
their natural enemies (Gonzlez-Hernndez et al. (Anderson 1926). The ant Pheidole punctata
1999). In the eld, Pheidole megacephala (F. Smith) was known to destroy the larvae and
(Fabricius) had a positive association with adults of Cryptolaemus montrouzieri Mulsant
Dysmicoccus neobrevipes Beardsley and a nega- preying on Planococcus citri in South Africa
tive association with the predators of mealybugs (Kirkpatrick 1927). The ineffectiveness of C.
(Jahn and Beardsley 2000). Collectively, P. mega- montrouzieri against Planococcus citri (Risso) at
cephala deters predators from attacking D. neo- Liguria (Italy) was due to the attack of ants such
brevipes. Ants are known to attack the parasitoids as Tapinoma erratium nigerrimum Oryl and
and predators of scales and mealybugs while Iridomyrmex humilis Mayr (Constantino 1935).
The presence of Argentine ant, I. humilis, Maconellicoccus hirsutus (Green) showed that
appeared to be partly responsible for the failure all C. montrouzieri introduced were killed and
of Cryptolaemus to become permanently estab- removed in 132.5 min. The mealybugs that asso-
lished in Bermuda (Bennett and Hughes 1959). ciated with ants are indeed protected from attack
In India, the failure of establishment of C. mon- by their predatory natural enemies, although
trouzieri in the mealybug-infested citrus orchards mealybugs and ants do not have an intimate asso-
of Assam was due to the activity of ant Oecophylla ciation (Lai YiChun and Chang NiannTai 2007).
smaragdina (Fab.) (Narayanan 1957). The con- There was an interaction involving the pink
trol of P. citri with Cryptolaemus was made inef- mealybug Sacchariococcus sacchari, the ant
fective in the presence of ant attendants (Panis Camponotus compressus (Fabricius), and the
and Brun 1971). Loss of results with C. mon- predator C. montrouzieri in sugarcane (Srikanth
trouzieri was caused by I. humilis in France et al. 2001). C. montrouzieri was found more in
(Greathead 1976). The ants Cremaster and numbers and proved successful against the
Iridomyrmex were known to prey on C. mon- mealybugs in the absence of ants (Van der Goot
trouzieri (Collins and Scott 1982). The failure of 1948; Murray 1982). In South Africa, the control
ant control is detrimental to biological control of of ants like I. humilis and Anoplolepis custodiens
citrus mealybug (Singh 1978; Narayanan 1957). F. aided the predator C. montrouzieri to give very
The observation of the protective behavior of P. good control of P. citri (Greathead 1971). Poutiers
megacephala against the attack of C. montrouz- (1922) also suggested protecting C. montrouzieri
ieri on the pink hibiscus mealybug from I. humilis in France.
15 Ant Association 201
Associations among invasive species of ants However, some mealybug predators, such as
and mealybugs are very important in their suc- coccinellids, apparently become tolerated by ants
cess in new locations (Helms and Vinson 2002). by mimicking the waxy body cover of the mealy-
The Argentine ant Linepithema humile (formerly bugs (Daane et al. 2007). This condition of para-
Iridomyrmex humilis) (Mayr) is an example of an sitepredator adaptation is often observed. The
invasive ant species that is a signicant pest in type of ladybird beetle larva is usually very mealy
both natural and managed habitats, and it is com- in appearance and blends in well with its pseudo-
monly associated with mealybug outbreaks coccid host. Thus, the parasite or predator spe-
(Daane et al. 2006; Silverman and Brightwell cies is well adapted to the use of ant-attended
2008). hosts. Ladybird beetle predators are often found
Saying that ants protect mealybugs from among formicid attendants. Ants seem unable to
natural enemies does not necessarily mean that recognize the ladybird beetle larva as a predator.
ants are attacking the natural enemies to save There is, therefore, little doubt that the seemingly
honeydew as a food resource. Ants deter the nat- mimetic resemblance of the beetle larva to the
ural enemies of mealybugs (Jahn and Beardsley mealybug is an aid to its more perfect predaceous
1994; Rohrbach et al. 1988). The encyrtid para- habitat. Another possible mealybug parasite
sitoid Anagyrus ananatis Gahan of pineapple adaptation has been noted in a species of y
mealybug is not only scared away from when larva. This larva apparently remains outside its
ants are present but they are also rarely killed by hosts body and extracts food externally. This
predators such as the ladybird beetles. When ants habit would seem to make it readily susceptible
are absent, the parasitoid is highly effective in to the attack by ants. In order to overcome the
lowering the mealybug populations in pineapple problem, the larva is always found completely
plantings. Ant and mealybug interactions were hidden beneath the mealybug, and only when the
studied in a pineapple eld near Honolu on the host insect is removed can the larva be seen.
island of Maui, Hawaii. Big-headed ant Pheidole
megacephala was found to have a positive asso-
ciation with gray pineapple mealybug 15.3 Ant Constructions
Dysmicoccus neobrevipes but no association
with Dysmicoccus brevipes (Cockerell). Sticky Ants protect the mealybugs from adverse weather
trap collections revealed that D. neobrevipes and by building earthen shelters around them and
D. brevipes are dispersed by the wind. The posi- moving them to protected places; and some ant
tive association between P. megacephala and D. species actively construct shelters for mealybugs
neobrevipes was not due to ants transporting that provide some protection from unfavorable
mealybugs but could have resulted from ants environments and natural enemies (Franco et al.
deterring natural enemies or removing honeydew 2000; Helms and Vinson 2002; McLeod et al.
(Gary and Beardsley 2000). 2002). There are two main types of ant construc-
202 M. Mani and C. Shivaraju
tions which are important to mealybugs. The rst drops of rain, although the tent itself is in no way
is the actual ants nest and the second is the so- waterproof. Second, the tent, which has only one
called carton or ant tent. The latter is not important very small entrance, is important in shielding the
in California, although Wheeler (1926) mentioned mealybugs from large parasites. Apparently, the
several instances of carton nests in North mealybug parasites are unable to either nd the
America. Ant-nest isolation of the mealybugs is tent entrance or push their way through. Although
considered quite important and has been observed there are many records of tent parasitism, the rate
on several occasions in California. In one instance, of incidence is much lower than where no protec-
Phenacoccus artemisiae Ehrhorn was found in tion is afforded.
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 15.4 Removal of Honeydew
instances were being transported in the mandibles
of an ant. This particular collection was made in Ants prevent the accumulation of honeydew by
early February, which is a time of very low insect consuming it (Jahn and Beardsley 1994; Rohrbach
activity and high precipitation rate. The mealybugs et al. 1988). Honeydew accumulation, and the
were quiescent for the most part and were found in sooty mold that can grow on honeydew, may be
close contact with one another on the ceiling of ant detrimental to mealybugs. These ants, especially
nest chambers. It seems likely also that parasite P. megacephala, have been blamed for protecting
protection would be important. them against their natural enemies while remov-
Fungus is also a parasite of mealybugs, ing the excess honeydew produced by the mealy-
although infestation is not normally observed bugs (Gonzlez-Hernndez et al. 1999).
until the pseudococcid has been mounted; when One of the direct benets of ant association to
examining mealybug preparations, however, fun- mealybugs is shown in relation to the production
gus infestation is often seen. Ant-nest protection of honeydew and the subsequent contamination
from a highly humid environment is therefore an of the honeydew and source insect with sooty
important factor in mealybug welfare and is prob- mold. Because this fungus contaminant is often
ably directly connected with protection from fun- the cause of mass destruction of mealybugs, cer-
gus contamination. Protection from harsh winter tain adaptations have been made to rid the mealy-
conditions is perhaps the most important factor of bug of the secretion. Ants remove honeydew
ant-nest benet to mealybugs in California. from mealybugs, thereby preventing fungi from
For the most part, ant tents are important in the attacking mealybugs, and the removal of honey-
tropical areas, where two primary types are found. dew prevents contamination, which may be espe-
The rst is constructed with the silk-forming cially detrimental to rst-instar nymphs (Cudjoe
glands of the ant larvae. The adult, which has no et al. 1993; Daane et al. 2006, 2007; Gullan and
silk gland, holds a larva in its mandibles and forces Kosztarab 1997; Moreno et al. 1987). Rohrbach
the immature form to produce its silken product in et al. (1988) hypothesized that honeydew feeding
the desired area. The ant genus Oecophylla is well by ants could benet mealybugs by preventing
known for this habit. The second type of tent is the accumulation of honeydew on the mealybugs
made of earth, paper formed by the ant, and themselves. Presumably, immature mealybugs
leaves. Any or all of these materials may play a get stuck in honeydew and die if ants do not
role in the tent construction. The tents are nor- remove it. Phenacoccus alieni McKenzie is
mally built over mealybug colonies which may known to squirt a small globule of honeydew
have as many as 1,200 individuals. Tent dimen- from the anus to a distance of over 4 in.. This
sions have been recorded up to 4.5 2.3 in. distance is at least 20 times the total length of the
Mealybug-derived benets are twofold. First, insect and shows that mealybugs are quite capa-
the tent provides some protection from direct ble of ejecting honeydew to distances well out of
15 Ant Association 203
the range of their personal contamination. When nest. Ants are the primary or sole means of
ants are in attendance, they remove the honeydew mealybug dispersal in pineapple. Illingworth
as described above, thus eliminating the problem (1931) observed P. megacephala carrying mealy-
of sooty-mold contamination. Because of this, bugs from one cage of pineapples to another. The
ant-attended mealybug colonies are quite often big-headed ant Pheidole megacephala. (F.),
very dense with little distance between individu- Argentine ant Linepithema humile (Mayr), and
als to allow for honeydew ejection. re ant Solenopsis geminata (F.) are commonly
However, in Californias coastal vineyards, found in the Hawaiian pineapple agroecosystem,
Argentine ants increased densities of the obscure where they tend pink pineapple mealybugs
mealybug Pseudococcus viburni (Signoret), pri- (PPM) and gray pineapple mealybugs (GPM) for
marily by removing the honeydew that impedes honeydew. These ants, especially P. megaceph-
the movement of crawlers. Meanwhile, the larvae ala, have been blamed for dispersing mealybugs
of C. montrouzieri successfully forage in patches (Gonzlez-Hernndez et al. 1999).
of high mealybug density. One hypothesis is that
larvae of C. montrouzieri, being also covered
with waxy structures, successfully mimic mealy- 15.6 Benet to Ants
bugs and avoid detection by ants. Furthermore,
when approached by an ant, the coccinellid larva Access to honeydew has been shown to enhance
stops moving and lowers its body against sub- the rate of increase of ant colonies. Honeydew
strate, thus better resembling a sessile mealybug. accounts for more than half of the diet of many
The ants move around the larva, and stroke it temperate wood ants (Formica spp.), and it is the
with their antennae like they stroke the mealy- dominant food source of some subterranean ants
bug. After failing to obtain the honeydew, the ant (Mittler and Douglas 2003). Mealybug exudates/
moves away. Densities of C. montrouzieri were honeydew is highly acidic (pH 3) with fructose
higher on ant-tended vines, where there were (45 g), glucose (20 %), and other sugar contents
more mealybugs (Daane et al. 2007). in negligible quantity (02 %) per 100 g of solids
Ants stimulated increased feeding by mealy- (Ashbolt and Inkerman 1990). The normal ant
bugs; tending by the ants may have other effects mealybug association when observed in the eld
that alter mealybug densities: It may also improve is seemingly quite simple. The ants, which may
the mealybugs habitat or tness (Daane et al. be of various genera, normally move busily from
2007). In the presence of ants, mealybugs are one mealybug specimen to another. When a
able to ingest larger quantities of sap (Degen and mealybug is contacted, the ant begins to fondle
Gersani 1989). In several instances, ant-tended the mealybug with its antennae just as it might
mealybug colonies will be much larger than colo- fondle its own brood.
nies of the same mealybug species on the same The ants rest their heads on the dorsum of the
host that are not tended by ants. Therefore, out- mealybugs near the area of the ostioles for hon-
wardly the ants presence must be of some benet eydew. Mealybugs extrude a solution from their
to the mealybug, either directly or indirectly. ostioles when disturbed. This is possibly a
defense mechanism. When Phenacoccus echeve-
ria McKenzie is purposefully disturbed, it
15.5 Transportation of Mealybugs extrudes two small globules of honeydew through
its posterior ostioles. An ant then comes along
Ants are known to transport the mealybugs from and within a few seconds ingests all of the
plant to plant between and within elds, thus extruded honeydew. It is clear that ants benet
facilitating mealybug dispersal. In California, it from mealybugs in receiving honeydew from
is often possible to see ant Camponotus actually them, which is added to the ants food supply.
carrying from its host plant, directly into the ants The amount that ants rely on honeydew for their
204 M. Mani and C. Shivaraju
existence varies greatly with the species involved. 15.7 Predatory Effect of Ants
Some ants seem almost entirely unknown
whether this adaptation to aid in procuring hon- A nal direct benet not often realized, but per-
eydew, or whether this is the normally in an adap- haps a factor of importance in the understanding
tation to aid in procuring honeydew or whether of the biological control of mealybugs, is that
the normally indigestible wax is actually used in some ants actively regulate the population size of
ants diet. their hosts. It has been demonstrated that certain
ant species will actually keep the mealybug pop-
ulation down to a size which they can control,
thus eliminating production of honeydew. The
ants regulate any excessive build-up of popula-
tion by killing a number of mealybugs, and con-
suming the mealybugs is considered a supplement
to the protein portion of the ant diet (Way 1963).
This habit is also of importance in eliminating
sooty-mold contamination in the mealybug col-
ony. Some ant species may switch between tend-
ing and preying on mealybugs (Degen and
Gersani 1989; Mittler and Douglas 2003; Way
1963). The predatory role of ants on the mealy-
bugs is very meager in the regulation of
mealybugs.
Globule of honeydew
A third and nal benet has been mentioned 15.8 Strange Aspects of Ant
earlier and deals with the use of associated Mealybug Associations
mealybug as a source of protein. Although preda-
ceous non-honeydew-consuming ants are not Exceedingly unusual observations were made by
known to attend mealybugs just for protein, some Bunzli in relation to an association between the
ants, which have both honeydew-consuming and mealybug Neorhizoecus coffeae (Laing) and the
predaceous habits, will occasionally kill mealy- ant Acropyga paramaribensis (Borgmeier). The
bugs and use them for protein. In nutritional eggs and immature mealybugs were kept in
value, honeydew is more complete than might be chambers with eggs and larvae of the ant. The
expected. It may contain free amino acids, immature female mealybugs apparently served as
amides, proteins, many minerals, and B vitamins honeydew sources. When the mealybug matured,
(Way 1963). The honeydew may vary greatly in it no longer produced honeydew and was then
its content depending on the species of mealy- transported by the worker ants to a separate
bug, the host plant, the age of plant, the part of mealybug chamber. The eggs that are laid by the
plant upon which mealybugs are feeding, and the mature females are then carried back to the brood
length of time that the insect feeds. Normally, a chambers. This association is not too unusual.
complete diet of honeydew will not compensate The extraordinary part of the relationship is the
for a protein deciency in the ant, and supple- fact that the winged virgin female ants, when
mentary protein must be added to correct the leaving the nest on their nuptial ight, always
situation. carry in their mandibles a fertilized female
There are several records of ants keeping mealybug. This mealybug will soon be the begin-
mealybugs pseudo wax-free. Ants might use ning of the honeydew source of a newly formed
mealybug wax for some nutritive value. Acropyga nest.
15 Ant Association 205
apple plants were applied with sticky bands an acceptable level in 23 years. The baits need
which had helped to prevent the movement of to be placed in the eld during budbreak (March
ants (Murray 1982). BHC solution (5 g/l) was to June, depending upon location in the state) at a
poured into the anthills prior to the release of C. rate of 1520 bait stations (UC bait station) per
montrouzieri against Ferrisia virgata (Cockerell) acre (Nyamukondiwa and Addison 2011). Ants
in guava orchards (Mani et al. 1990). In the can also be managed by applying tanglefoot
orchards where ants were partially excluded, a every 12 weeks.
signicant reduction in citrus mealybug popula-
tions and damage could be observed (Villalba
et al. 2006). Applying a 6 % solution of chlorpy- References
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Constantino G (1935) Unnemicodel contonela degli-
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Gourmet ant bait (Innovative Pest Control Pseudococcidae) in Ghana. Bull Entomol Res
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insecticide baits on Argentine ants in Californias
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Linepithema humile (Mayr), common pugna- Daane KM, Sime KR, Fallon J, Cooper ML (2007)
cious ants Anoplolepis custodiens (F. Smith) and Impacts of Argentine ants on mealybugs and their
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Argentine ants is important in mealybug manage-
(Homoptera, Pseudococcidae). J Zool 218:421432
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11(2):167170 the biological control of citrus and coffee mealybugs.
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64:12 producing Homoptera. Annu Rev Entomol 8:307344
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Inuence of inoculative releases of natural enemies and behaviour. Columbia University Press, New York,
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citrus mealybug Planococcus citri (Hemiptera:
Methods of Control
16
M. Mani and C. Shivaraju
Mealybugs pose a serious threat to several agri- ior of mealybugs may provide a spatial refuge
cultural and horticultural crops throughout the from natural enemies and harsh environmental
world. They are a major problem in greenhouses conditions (Berlinger and Golberg 1978;
and nurseries, where they often cause severe eco- Gutierrez et al. 2008a). This type of plant coloni-
nomic damage. It seems surprising that such a zation makes mealybugs practically invisible
delicate soft-bodied insect is so difcult to con- during the latent population phase. However, dur-
trol. There are, however, several reasons which ing outbreaks, the population boils over from
may account for this fact. the refuge and becomes conspicuous. In addition,
Mealybugs developed several different other species have the habit of spending their
defense mechanisms. Many of the species tend to entire lives deep in the soil, protected almost
establish themselves in protected sites, such as from insecticidal materials.
cracks and crevices in bark, leaf axils, root Mealybugs are noted for the production of
crowns, nodes of grass stems, under fruit sepals dermal wax secretions. Adult mealybugs and the
and within fruit navels, between touching fruits nymphal instars are covered with waxy coating.
or fruits and leafs, and in tunnels bored by insect Also the eggs of mealybugs, protected by the
larvae in roots and stems (Franco et al. 2000; waxy lamentous secretions of the ovisac, are
Kosztarab and Kozr 1988). This cryptic behav- almost impossible to reach with insecticides.
Eggs protected with waxy ovisac Adult mealybug covered with Mealybug crawlers not covered
waxy coating with waxy coating
The waxy secretion is the most common con- control of some mealybugs. The purpose of this
spicuous trait of the mealybug family. It is a com- study is not to give specic recommendations for
plex system that serves different functions, and mealybug control. It will be dealt under the
which is produced by the epidermal wax glands respective crops. General methods of mealybug
and transported to the body surface via ducts, control are summarized in this chapter.
pores, and secretory setae of various types (Foldi
1983; Gullan and Kosztarab 1997). The main
components of the wax of ve mealybug species 16.1 Decision-Making System
(Planococcus citri (Risso), Pl. cus (Signoret),
Pl. vovae, Pseudococcus cryptus Hempel, and The ve key elements ought to be considered in a
Nipaecoccus viridis (Newstead)) were trialkyl decision-making system in the management of
glycerols and wax esters. The wax cover is mealybug population are (1) information on
believed to prevent water loss. The hydrophobic mealybug density, perhaps obtained late in the
property of the wax enables the mealybugs to season, in case of overwintering population; (2)
escape drowning or becoming swamped by water awareness of the population distribution in the
in their typical cryptic sites. The ovisac, which is target area; (3) information about the density of
also a wax secretion, is considered to be an adap- the relevant natural enemies; (4) the density of
tation that protects the offspring from both wet associated insect species, which may increase the
and dry conditions, and that may also provide an damage or render the activity of the natural ene-
attachment to the host plant. Tubular ducts and mies; and (5) the risk of the spread of mealybug-
multilocular disc pores, respectively, produce transmitted viral disease. In this chapter, the
long hollow and shorter curled laments, which current knowledge needed to take actions and to
make up the ovisac and the male cocoon (Cox suggest solutions for different situations is
and Pearce 1983; Foldi 1983). The white wax of reviewed. Based on such knowledge, the grower
mealybugs is strongly light reective and may may select the appropriate control tactics.
reduce desiccation. In some cases, the wax also
serves to cover the honeydew droplets and to pro-
tect the mealybugs from contamination by their 16.2 Monitoring
own honeydew and defensive exudates (Gullan
and Kosztarab 1997). Normally, chemicals are Vigilance is important to eliminate them before
used to control the mealybugs. However, the there is a major outbreak of mealybugs.
crawler stage is not covered with wax, and as a Monitoring for the incidence of mealybugs is a
consequence, this is perhaps one of the most sus- prerequisite to initiate the management practices.
ceptible stages of mealybug to chemicals. On the There are no simple and effective methods to
other hand, natural enemies have given excellent visually monitor the mealybugs, and the process
16 Methods of Control 211
itself can be time consuming and laborious. As and, therefore, the most effective timing for the
exemplied for Pseudococcus maritimus, the management options.
accuracy of monitoring the plant material will
depend on the mealybug population density, and
the number of samples needed for an accurate 16.3 Cultural Method
count is often high because most of the mealy-
bugs have a clumped distribution pattern, often A number of cultural controls are practiced and
being found on only a small percentage of the these vary greatly among regions. Crop sanita-
plants. The appropriate sampling program will tion is useful in reducing the mealybug popula-
also vary throughout the season, depending tion. Before applying the insecticides/pesticides,
largely on mealybug location as there are periods manual removal of the uffy nests and most of
when much of the population is hidden (e.g., the insects is advisable. It greatly increases the
under bark) rather than exposed (e.g., on leaves). chance on complete elimination of the mealy-
In addition, as species have different numbers of bugs. Removal of the weeds harboring the mealy-
annual generations and preferred feeding loca- bugs eliminates the source of mealybug
tions throughout the season, there is not a single infestation. For example, crop sanitation includ-
sampling procedure appropriate for all the mealy- ing the removal of weeds was useful in the con-
bugs. In most cases, signals of an infested plant trol of Heterococcus pulverarius (Newstead)
can be used to aid the sampling program. First, (Dietz and Harwood 1960) and H. nigriensis
ants are closely associated with the mealybugs, Williams (Harris 1961). Burning and plowing the
and their presence can help select the plants for crop after harvest result in very little reoccur-
further sampling. Second, honeydew on the rence of the mealybugs. The control of certain
leaves can also be a good signal; a large popula- garden mealybugs may be done simply by host-
tion hidden under the bark will excrete enough ing the plants down with a strong stream of water
honeydew that the infested trunk region will have (Michelbacher et al. 1959). Although this seems
a darker, wet appearance. Third, as mealybug rather unorthodox, the control is fairly success-
numbers build up, their feeding damage may ful, especially if this treatment is used at regular
cause leaves to turn yellow or brown and drop intervals. In the case of woody plants, mealybugs
from the plant. Finally, during the harvest time, are found underneath the bark of the trunk, cor-
fruits in direct contact with the spurs or trunk are don, spurs, and canes. These locations provide
more likely to be infested, and by selecting these some protection from insecticides, natural ene-
fruits, a higher mealybug count can be made A mies, and environmental conditions. Stripping
faster sampling method is to use sticky traps the bark exposes the mealybugs to these mortal-
baited with sex pheromone to lure in and trap the ity factors. The infested bark should be destroyed
adult winged males. It has long been known that rather than left in the row middles as the mealy-
sexually mature female mealybugs like bugs can move back. Cover crops have been used
Planococcus citri emit a sex pheromone to attract to improve soil health and lower-pest densities by
the winged adult males. These pheromones can increasing the natural enemy numbers or
be synthesized and used in the eld for monitor- diversity.
ing the mealybugs. Thus, the monitoring provides Parasitoids that attack the mealybugs could
essential information for making decisions indi- utilize oral nectaries of some cover-crop species
cating the presence and the numbers of mealy- as a food source to increase adult longevity.
bugs and their enemies occurring in nature; the Generalist predators, such as the lacewings and
degree of natural control by insects, which prey some ladybird beetle species, might also utilize
upon or parasitize the mealybugs; whether the these oral food resources as well as herbivores
mealybugs are high enough to require treatment; in the cover crop as alternate prey. Overly vigor-
the lifecycle stages of any mealybugs present ous plants can increase mealybug populations in
212 M. Mani and C. Shivaraju
two ways. First, excess nitrogen has been shown mealybugs because the mealybug waxy covering
to increase the size of mealybug females and the repels polar chemicals (Walton et al. 2004).
number of eggs in each ovisac. Second, the Insecticides, with contact and also systemic
increased foliage associated with overly vigorous activity, are still primarily used to control or reg-
plants provides better shelter for the mealybugs ulate the mealybug populations.
by reducing temperatures inside the vine leaf Normally chemicals are used to control the
canopy, and may reduce the amount of applied mealybugs. There are great similarities among
foliar insecticide that reaches the mealybug. the insecticide arsenals used to control mealybug
Controlling plant vigor is, therefore, a practice species on different crops. In principle, three
that can help improve mealybug control. main modes of insecticide application are
adopted: (1) foliage cover spraying for manage-
ment of aboveground populations; (2) applica-
16.4 Physical Control tion of insecticide solution to the soil to enable it
to penetrate to the root zone, so as to combat sub-
The following physical control measures can be terranean colonies; and (3) chemigation by appli-
adopted to reduce the mealybug population on cation of systemic compounds via the irrigation
the plants: (1) ushing mealybugs off the leaves system (Chemigation), for example, drip irriga-
with water can provide immediate relief but will tion. Insecticides are also used against mealybugs
simply displace the mealybugs; (2) rubbing off by smearing them on the stem or main branches.
and crushing the colonies with a cloth; (3) mealy- For example, swabbing of grapevine trunk/stem
bugs can also be removed by dipping a cotton with chlorpyriphos is recommended to control
swab in alcohol or ngernail polish remover; (4) the mealybugs. Two other, less common, tech-
discarding heavily infested plants; and (5) prun- niques are fumigation, usually applied for eradi-
ing infested tissue off infested plants. cation, for example, with methyl bromide, and
slow-release strips to prevent colonization.
pests does not exist in practice. Since the growers organophosphates such as malathion, diazinon,
will need to treat small hot spots of the mealybug, tetraethyl pyrophosphate (TEPP), and dimefox
it is expected that some soft insecticides will be have been used with partial success but they are
used and that more than one application may be not in use now due to one reason or the other.
needed to selectively eliminate such hot spots. They may be extremely hazardous or can develop
When these hot spots are treated, several points resistance (Madsen and Westgard 1962).
should be taken into account: (1) the hot spots are Malathion is primarily used in the control of gar-
expected to be in areas that are practically free of den and nursery mealybugs (Michelbacher et al.
problematic mealybug populations; they actually 1959). TEPP, another organophosphate, has
constitute oases for parasitoids and predators; effectively controlled P. citri, Ps. longispinus,
therefore, the ratio of mealybug to natural enemy and Ps. maritimus (Jefferson and Pritchard 1961).
populations in the hot spots should be considered Some old organophosphates, such as dichlorvos
before initiation of any control operation; (2) an and chlorpyriphos, are still being used against
insecticide will be applied when augmentation mealybugs because they certainly are much less
with predators is not useful or cannot be imple- dangerous. Many of the chlorinated hydrocar-
mented; (3) a low-residue short-life insecticide is bons, such as dichlorodiphenyltrichloroethane
the most appropriate; (4) an augmentation of (DDT), lindane, aldrin, dieldrin, and endrin, and
natural enemies will be needed if the hot spots the organophosphate (parathion) are not in use
are too numerous. now due to various reasons. Eventually, however,
most of these materials became less effective
16.5.1.1 Application Timing also. Organophosphates, such as chlorpyrifos,
Application timing is critical to control mealy- acephate, dichlorvos, and diazinon, and, to a
bugs with most insecticides. Exposed mealybugs lesser extent, carbamates, such as aminocarb,
are more easily killed than those under the bark, carbaryl, thiodicarb, or methomyl, are broad-
and the smaller stages are more susceptible than spectrum nerve insecticides, which have been
the larger mealybugs. This is especially true for used against mealybugs that colonize the plant
insecticides with a short residual period. Much canopy since the early 1960s (Gonzalez et al.
research, therefore, has been aimed at proper 2001; Shafqat et al. 2007). These insecticides
application timing and development of materials when applied in high volume could successfully
with better penetration into the protected habitats overcome the obstacles that make mealybugs
of mealybugs. Applications with systemic insec- hard to kill. The obstacles are as follows: (1) their
ticides near bloom are often used, as the insecti- hydrophobic wax cover, which repels hydrophilic
cide moves quickly in the plants. insecticides; (2) their tendency to feed in hidden
and protected parts of the plant; (3) their typically
dense colonies; and (4) the frequent overlapping
16.5.2 Foliar Spray of generations. Effective control is achieved
when most of the mealybug population is in the
The dispersive habit of the crawler should make dispersive crawler stage or the young nymphal
it more susceptible to insecticides than the later instars, and when the host plant does not provide
developmental stages that live in sheltered sites. effective shelter. However, satisfactory control is
Spray application is to be timed to coincide with often difcult to achieve over an extended period.
the crawler stage as it would be effective and These chemicals have detrimental effects on the
would also permit the use of less persistent environment as a whole and on natural enemies
chemicals. in particular (Anand and Ayub 2000; Babu and
Not many specic insecticides are available Ramanamurthy 1998; Meyerdirk et al. 1982).
against all the species of mealybugs, but para- The multivoltine character of the pest mealybugs
thion was primarily used as spray and dust in and the frequent application of inefcient control
commercial agriculture. In the earlier years, measures accelerate the development of insecti-
214 M. Mani and C. Shivaraju
cide resistance (Flaherty et al. 1982). Systemic and thereby improve the insecticide efcacy
organophosphates such as dimethoate could (Cranshaw et al. 2000; Morishita 2005). Summer
overcome some of these obstacles (Grout and oil emulsions/mineral oils are particularly effec-
Stephen 2005; Meyerdirk et al. 1982; Prasad tive in the control of mealybugs on ornamental
et al. 1998). Chlorpyrifos-impregnated strips are plants. Applications should be made at regular
applied to protect banana bunches from the intervals of 13 weeks (Michelbacher et al.
mealybug infestation or applied as stem barriers 1959). Combinations of these oil emulsions with
for the control of ants (Addison 2002; Gross et al. contact insecticides are quite effective in the con-
2001). trol of garden and household mealybugs. Neem
Newer materials, with more novel modes of oil, horticultural oil, and insecticidal soaps are
action, have also gained in popularity, including often regarded as organic or non-chemical
neonicotinoids, insect growth regulators (IGRs), methods, but this is not completely accurate.
botanicals, and biosynthesis inhibitors. However, they are safer than the insecticides.
Application of spirotetramat 6 . oz/acre + adju- They will not provide absolute control over
vant (Ventre 0.25 % v/v) 44 . oz/acre at a spray mealybugs but can drastically reduce their popu-
volume of 137 gal/ac was able to reduce the lations. ChilliGarlic extract is also used to con-
bunch infestation with mealybugs to 3 % fruit trol the mealybugs.
damage and there was little to no honeydew in
that treatment. A major difference between the Botanicals
older and newer materials is the importance of Neem has natural insecticidal properties but is
coverage. As mentioned, a portion of the mealy- biodegradable and non-toxic to several naturally
bug population is often under the bark and, for occurring parasitoids and predators. It works by
some species, on the roots. Many of the older making the leaves unpalatable to the mealybugs.
foliar sprays did not effectively contact and kill Neem is to be sprayed like other contact insecti-
mealybugs in these more protected locations. cides. Spraying should be in such a way that the
Some of the more novel materials have systemic undersides of all leaves are covered. In organic
properties, applied either through the irrigation agriculture, azadirachtin, an IGR chitin inhibitor
system or as a foliar spray. For organic or sustain- derived from the Indian neem tree, may be used
able farming programs, neem, light mineral oils, in similar modes (Irulandi et al. 2001). Pyrethrins
lime sulfur, citrus products, and fatty-acid soaps and rotenone replaced some of the old com-
have been used. pounds in organic agriculture with limited effec-
Another historical difference is that the earlier tiveness. Neem products have a repellent effect
materials were often broad spectrum and killed on some mealybugs. Neem oil is effective for
more than just the targeted mealybugs. The mealybug suppression. Neem oil is generally
extensive use of DDT and other synthetic insecti- considered safe for humans, pets, and plants
cides to control leafhoppers apparently disrupted unlike usual chemical insecticides. Neem oil is
the natural control of the mealybug P. an all-natural organic insecticide. Unlike the
maritimus. toxic chemicals, neem oil interrupts the pest
reproduction cycle and is, therefore, useful in
16.5.2.1 Oil Emulsions/Mineral Oils/ eliminating mealybugs from the plants. Mix
Botanicals 5 ml (1 oz) of pure neem oil with 2.5 ml (1/2 oz)
of a mild liquid soap and 1,000 ml of water (four
Oils cups). Mix neem oil and liquid soap rst and
Oils have long been used for the control of scale then add water. Mix it thoroughly and spray.
insects but they have been ineffective against Neem oil solution smothers the mealybugs;
mealybugs. However, the integration of narrow therefore, complete coverage of all the plant
rened oils with other insecticides was suggested parts is essential. Repeat every 57 days until the
as a means to dissolve the insects wax covering infestation comes under control. Bug Buster is a
16 Methods of Control 215
botanical solution for mealybugs. Two sprays of Dishwashing soap can be used as an effective
Bug Buster are recommended; rst, at a dose of remedy for mealybugs. According to evergrow-
45 ml/l of water at an interval of 35 days and ing.com, combine one tablespoon of dish soap
thereafter, at an interval of 715 days. Bug with one pint of warm water. Mix the solution in
Buster acts as a contact as well as systemic a spray bottle and coat the plants with a layer of
against mealybugs; it penetrates the waxy cover the solution. The soap penetrates the protective
of the insects body and eventually kills them; waxy coat created by mealybugs and kills the
and it disrupts the structure and permeability of pests. Check all areas of the plant, including the
the insect cell membranes. Disruption of cell underside regions, for infestation. Spray every
wall damages the cells resulting in quick killing region of the plant to ensure complete
of the mealybugs. Hence, Bug Buster is very eradication.
effective for all the stages of mealybug, as it is
biodegradable and safe for humans; compatible
with most of the bio-pesticides/-fertilizers; free 16.5.3 Soil Drench
from harmful synthetic chemicals and water sol-
uble; acts as systemic as well as contact against Soil drenching with malathion and parathion
mealybugs; non-hazardous; safe to humans and were partially effective against root mealybugs.
pests; and non-polluting, eco-friendly, and no Heavy infestations of Rhizococcus pritchardi
residual toxicity. Because it is a combination of McKenzie on roots of African violet have been
active natural extracts, there is no possibility of successfully controlled by drenching the potted
developing resistance (http://www.ehow.com/ plants dimethoate (Snetsinger 1966). At present,
list_7578648_home- remedies- mealybugs. the mealybug management is based on chemical
html#ixzz2sd1mn MBI). Unripe fruit extract of treatments, primarily with neonicotinoid insecti-
the plant Balanites aegyptiaca showed inhibition cides (e.g., imidacloprid, thiamethoxam, clo-
of the mealybug Ferrisia virgata (Cockerell) thianidin). These are typically applied as a soil
after the third day of spraying. No mealybugs drench directed to the roots. Soil drench applica-
were observed on the leaf on the seventh day of tions of imidacloprid is highly effective in reduc-
the application (Wabale et al. 2010). About 90 % ing the mealybug populations, particularly when
mortality of Planococcus citri was obtained with applied at 0.525-g ai/vine makes it extremely
pepper and eucalyptus extract at 3,500 ppm effective. In California, the mealybugs were also
(Ahmadi et al. 2012). controlled when imidacloprid was applied
through irrigation lines or into furrows (Sazo
Horticultural Oil et al. 2006). A further benet of soil drenching is
Horticultural oils are petroleum distillates. They that the insecticide is transported to the pest with-
are to be applied underneath leaves, on pots, and out harming the predators and parasitoids and
areas surrounding the plants. These oils (if not can be applied before they are active. Under this
phytotoxic) should not be applied to plants when condition, imidacloprid could be used to kill
temperature is greater than 85 F or in direct mealybugs on the roots of the plants. Imidacloprid
sunlight. soil treatments have good residual activity and
the control is sustained even up to 2 years.
Insecticidal Soaps Systemic insecticides are applied preventa-
Insecticidal soaps are a solution of synthetic tively to the growing medium as a drench or as a
pyrethroids mixed with a mild detergent made granule for uptake or absorption via the roots and
from petroleum products. These soaps (if not then translocated throughout the plant through
phytotoxic) should be applied underneath leaves, the vascular system. Most systemic insecticides
on pots, and areas surrounding the plants and are translocated through the plant via the transpi-
should also be used on greenhouse vegetables. ration stream, which is the movement of water
216 M. Mani and C. Shivaraju
through the plant by means of the xylem- or in the best control of Planococcus citri (Risso)
water-conducting tissues. They are primarily infesting the gardens (Bivins and Deal 1973).
active on phloem-feeding insect pests with pierc-
ingsucking mouthparts, such as mealybugs, as
these insect pests feed exclusively within the 16.5.4 Insect Growth Regulators
xylem vessel elements or phloem sieve tubes.
During the feeding process, these insects with- The chemicals having the IGR activity are used
draw and ingest lethal concentrations of the sys- to reduce the mealybug population. For example,
temic insecticides active ingredient and are the IGR ZR-777 (prop-2-ynyl 3,7,11-trimethyl-
subsequently killed. There are a number of (2E,4E)-dodecadienoate), gave good control of
advantages associated with using drench or gran- nymphs in all instars of Planococcus citri (Risso)
ular applications of systemic insecticides com- receiving 0.01 % sprays. One foliar application
pared to foliar sprays. For instance, drench of ZR-777 gave good control of Pseudococcus
applications reduce exposure to workers and longispinus (Targ.) and Phenacoccus solani
natural enemies, such as parasitoids and preda- Ferris after 5 days of application. The IGRs, such
tors. In addition, systemic insecticides are trans- as buprofezin, a chitin-synthesis inhibitor, or kin-
located through the plant vascular system oprene, which mimics juvenile hormone, were
including the xylem and phloem, protecting the sought as replacements for organophosphates
growth that would have been missed when apply- and carbamates in controlling mealybugs; they
ing a contact insecticide, as well as any new have been considered a suitable alternative
growth following the application. This may pro- because they exhibit low human toxicity. They
vide protection for extended periods of time. are more selective to many benecial species and
Furthermore, applying systemic insecticides as they are specically targeted at processes
drenches reduces the amount of material lost due involved in particular stages of mealybug devel-
to evaporation, light degradation, and irrigation opment. However, many of the IGRs are toxic to
(wash-off). ladybeetles (James 2004; Cloyd and Dickinson
2006). Buprofezin is a commonly applied IGR
16.5.3.1 Soil Application against mealybugs (Muthukrishnan et al. 2005);
Soil application of granular insecticides, namely however, its effectiveness is mainly limited to
phorate, disyston, and aldicarb, has been recom- eggs and young stages, so that the adult females
mended to control the mealybugs. Phorate has may escape the consequences of the treatment.
been used in the control of Heterococcus pulver- Buprofezin also suffers from the same limitations
arious (Dietz and Harwood 1960), Ferrisia vir- as other foliar-sprayed compounds. Buprofezin
gata (Ckll.), Planococcoides njalensis (Laing), (Applaud) is emerging as a prime control tool for
and Ps. comstocki (Kuwana) (Abrahao and mealybugs. When applied, for three seasons, the
Mamprim 1958). Disyston has been used in the chemical works extremely well for the mealy-
control of Dysmicoccus brevipes (Ckll) (Carter bugs. It is an IGR, and great control is achieved
and Gortner 1958). while allowing benecial insects to continue
Aldicarb granules applied in soil resulted in feeding with no notable disruption. Chlorpyriphos
excellent control of Maconellicoccus hirsutus is by far the most popular pre-harvest material,
(Green) (Mani and Thontadarya 1991). In but buprofezin can also be applied post harvest.
Shanghai region, aldicarb (5 %) granules mixed The IGR Applaud provides effective control of
in soil around the ornamental succulent other sucking pests like soft scales, ash whitey,
Kalanchoe blossfeldiana plant roots or 40 % etc. The active ingredient, buprofezin, is
omethoate 100 poured over the roots resulted in extremely effective against crawler and nymph
97 % control of Planococcus citri (Tang et al. stages of these pests by inhibiting chitin biosyn-
1992). Aldicarb when applied as 10 % granules thesis. Application of the juvenile hormone ana-
into the soil with a drench of l water/plant resulted logue epofenonane (RO 103108) at 10 ppm,
16 Methods of Control 217
together with adjuvants, inhibited the male and impacts of the semiochemicals are limited to the
female development of Pseudococcus calceolar- target pest organisms. The potential of mealybug
iae (Mask.) (Rotundo 1978). sex pheromones as an alternative and ecologi-
cally friendly means for monitoring and control
16.5.4.1 Pesticides Known to Control is important and promising. Sex pheromones are
Mealybugs used in lures for monitoring, for detection of out-
Pesticides that are known to control mealybugs breaks, and for population management.
are as follows: acephate, acetamiprid, azadi- Monitoring systems provides vital information
rachtin, bendiocarb, bifenthrin, buprofezin, car- for the timing of insecticide applications.
baryl, chlorpyrifos, clothianidin, Population levels can be reduced or controlled by
cyuthrin + chlorpyrifos, cyuthrin + imidaclo- mass trapping, mating disruption, or lure and kill.
prid, diazinon, dimethoate, fenpropathrin, onic- The success of these methods depends on the
amid, uvalinate, imidacloprid, kinoprene, availability of the pheromone and on an appropri-
lambda-cyhalothrin, malathion, permethrin, pyri- ate formulation and deployment. In contrast to
proxyfen, S-kinoprene, and thiamethoxam. the extensive use of sex pheromones in control-
ling the beetle and moth pests, sex pheromones
16.5.4.2 Precautions are not yet employed in the control operation of
Pesticides can provide short-term control but are scale insects (Franco et al. 2009).
not recommended for long-term control because
mealybugs often persist in hard-to-reach areas.
Mealybugs are most susceptible to chemicals 16.6 Biological Control
when they are in the crawler stage. A waiting
period of at least 2 weeks after using pesticides Although it is apparent that many problems arise
before releasing the biological control agents is when chemicals are applied for the control of any
needed. High-volume wet sprays are needed in insects, some of these aftereffects warrant special
order to penetrate the waxy coating that protects mention in the case of mealybug control. Often
mealybugs. It may take a series of applications at when applications of insecticides are made for
10- to 14-day intervals to control mealybugs. the control of certain mealybugs, instead of caus-
Repeated use of the same pesticide or the pesti- ing a decrease in the mealybug population, an
cide combination more than three times in a row outbreak is also noted. The density of a Japanese
should be avoided. mealybug, Planococcus kraunhiae, on Japanese
persimmon fruit was higher in plots frequently
16.5.4.3 Pheromone-Based treated with cypermethrin than that in the
Management Tactics untreated plot. The number of mealybugs found
Sex pheromones of insects, including mealybugs, on Fuyu, a non-astringent cultivar, was higher
are natural compounds emitted by virgin females than that on Hiratanenashi, an astringent culti-
in order to attract conspecic males for mating. It var (Morishita 2005). This is usually because the
has long been known that sexually mature female insecticides have little effect on the mealybug,
Planococcus citri emit a sex pheromone to attract but eliminate the natural enemies, which were at
the winged adult males. These pheromones can one time holding the mealybug population in
be synthesized and used in the eld. Several check. At times, certain insecticides control the
chemicals have been identied to attract the mealybugs; however, with the decrease of com-
mealybugs like P. citri, Maconellicoccus hirsu- petition, other pests, particularly mites, increase.
tus, etc. The sex pheromones are effective in This second pest may become a more serious pest
extremely small quantities; they are non-toxic than the rst. Finally, it has often been noted that,
and can be applied in various ways. Unlike pesti- although a chemical may give excellent control
cides, these chemicals are species specic and do for a short time, the mealybug population, when
not affect benecial insects. The behavioral building up again, will reach an even higher level
218 M. Mani and C. Shivaraju
than it had attained before the chemicals were mealybugs and conrmed to be pathogenic; they
rst applied. The frequent use of insecticides and include Aspergillus parasiticus Speare,
labor for mealybug control has made their cost to Cladosporium oxysporum (Berk and Curt.),
the grower still greater. Hirsutella sphaerospora H.C. Evans and Samson,
Mealybugs have many natural enemies, and Neozygites fumosa (Speare) Remaudire and
including parasitic wasps, arthropod predators, Keller (Browning 1994; Delalibera et al. 1997;
and entomopathogenic fungi. However, parasit- Le Ru 1986; Moore 1988; Samways and Grech
oid encyrtids and predatory ladybird beetles 1986).
(Coccinellidae) are the most common natural
enemies of mealybugs, and a tremendous amount
of research has been done in this area, much of it 16.6.1 Classical Biological Control
quite successfully.
Mealybug-parasitizing encyrtids are primary Biological control of mealybugs has been prac-
endoparasitoids, most of them undergo solitary ticed for many years; it involves three main tac-
development. Coccidoxenoides, Gyranusoidea, tics, that is, classical biological control,
Leptomastidea, Leptomastix, Pseudaphycus, and augmentative releases, and conservation biologi-
Tetracnemoidea are examples of encyrtid genera cal control. Species are considered invasive if
of mealybug parasitoids (Charles 1993; Franco they are transported outside their native range
et al. 2000; Noyes and Hayat 1994; Rosen 1981). and become established, spread, and adversely
A number of predators contribute to mealybug affect the environment. Since the mealybugs are
control. Few specialize on mealybugs, whereas the most invasive species, classical biological
most are generalists that prey on any small, soft- control has been frequently employed against
bodied arthropods. Sometimes, the naturally them. Moore (1988) reviewed the natural ene-
occurring parasitoids, including encyrtids and mies used against mealybugs in biological con-
aphelinids, and predators, such as coccinellids, trol programs worldwide. According to Moore,
lacewings, cecidomyiids, and drosophilids, play more than 70 species of parasitoids have been
a major role in the suppression of the mealybugs. introduced against mealybugs, and at least 16 %
Invasive mealybugs are often being controlled of the introduced parasitoids were considered to
excellently with the introduced parasitoids and initiate substantial or complete control. Most of
predators. Coccinellids accept a wide range of the introduced parasitoid species were encyrtids,
food, but they complete the larval development but species of Aphelinidae and Platygastridae
and produce viable progeny only if they consume proved to be successful on several occasions.
their essential food. Four genera of Chilocorinae Often a single parasitoid was considered to be
(Brumus, Aspidimerus, Stictobura, and Orcus) responsible for the success, even when more than
and six genera of Scymninae (Diomus, Nephus, one was introduced. Noyes and Hayat (1994)
Sidis, Parasidis, Cryptolaemus, and reviewed the use of encyrtids for biological con-
Pseudoscymnus) prey preferentially on mealy- trol of pest mealybugs and found that out of a
bugs (Iperti 1999). Other important groups of total of 385 importations of encyrtids, targeting
predators are brown lacewings (Neuroptera; 22 mealybug species, about 24 and 7 % were
Hemerobiidae) and predatory gall midges considered to give partial or successful control in
(Diptera; Cecidomyiidae). The most well-known the eld and in greenhouses, respectively. With
predator is the mealybug destroyer, Cryptolaemus regard to predators, Moore (1988) analyzed the
montrouzieri Mulsant, which is native to use of C. montrouzieri separately from that of
Australia. other mealybug predators. This ladybeetle has
As sap feeders, mealybugs are not likely to be been used many times against at least ten differ-
exposed to viral or bacterial infections (Moore ent species of mealybugs and was considered to
1988) and only a few species of entomopatho- give substantial or partial control in about 19 %
genic fungi were reported to be associated with of the introduced predators; on some occasions, it
16 Methods of Control 219
has been regarded as an outstanding biological (Bokonon-Ganta et al. 2002), the pink hibiscus
control success. Of the other 46 predator species, mealybug, Maconellicoccus hirsutus (Green) in
mostly coccinellids, as well as cecidomyiids, the Caribbean and California (Roltsch et al.
chrysopids, hemerobiids, and lycaenids used in 2006), and the papaya mealybug, Paracoccus
biological control of mealybugs, only the cecido- marginatus Williams and Granara de Willink in
myiid, Kalodiplosis pseudococci Felt, was Palau (Muniappan et al. 2006). It is important to
regarded as having given signicant control, note that successes were mostly achieved in trop-
when used against Dysmicoccus brevipes ical regions where the target area for classical
(Cockerell) in Hawaii in conjunction with two biological control and the area of origin of the
parasitoids. Stiling (1993) showed that the major introduced parasitoids displayed similar climatic
reason for the failure of introduced natural ene- conditions. In a few cases, modeling has been
mies to reduce the pest population is related to used as a tool to analyze actual systems and to
climate (34.5 %). Moore (1988) analyzed the rea- identify major constraints, in order to improve
sons for the failure of both parasitoids and preda- the biological control of mealybugs. For exam-
tors of mealybugs to become established in ple, the model developed by Gutierrez et al.
biological control programs. In the case of para- (2008a) predicted that the parasitoid A. pseudo-
sitoids, Moore cites the following documented cocci would have a larger impact on the vine
reasons: (1) incorrect identication of the target mealybug P. cus than either L. abnormis or C.
mealybug species; (2) the target was a native spe- montrouzieri, and that biological control of the
cies; (3) hyperparasitism; (4) failure of the para- mealybug in California would require additional
sitoid to adapt to unfavorable climates; and (5) species of natural enemies and/or could be
other reasons, such as interference with ants, use achieved by reducing the size of the spatial-
of pesticides, and small numbers of individuals temporal refuge. In another use of a modeling
released. With regard to predators, Moore (1988) approach, Gutierrez et al. (2008b) concluded that
listed six main reasons for failure: (1) no adapta- the biological control of the vine mealybug might
tion of the released species to climate; (2) effect be adversely affected by climate change.
of the pesticides; (3) density of the prey; (4) Gutierrez et al. (1993) developed a tritrophic
effect of the host plant; (5) inability to reach the model of the cassava system and used it to explore
prey; and (6) effects of the other organisms. The the basis for the successful control of the cassava
lack of adequate food resources for natural ene- mealybug P. manihoti in Africa by the exotic
mies within or near to agroecosystems may limit parasitoid Epidinocarsis lopezi (DeSantis), and
the performance of biological control agents also to examine the causes for the failure of the
against mealybugs. For example, Davies et al. related parasitoid E. diversicornis (Howard) to
(2004) observed that the survival and reproduc- establish itself.
tion of Coccidoxenoides perminutus Girault, a
parasitoid of the citrus mealybug Pl. citri, were
signicantly inuenced by the nature of the nec- 16.6.2 Augmentative Control Tactics
tar on which the parasitoid was fed. In light of
these results, it was suggested that the habitat The rst known case of an augmentative biologi-
management, for example, by providing suitable cal control program dates back to before 1917 and
nectar sources for adult parasitoids, might be a was aimed at controlling the citrophilus mealybug
means to conserve and enhance C. perminutus Ps. calceolariae, a pest of citrus in Southern
activity in the eld. In recent years, successful California, by using the coccinellid predator C.
classical biological control programs against montrouzieri (Luck and Forster 2003; van
mealybugs have targeted the cassava mealybug, Lenteren 2006). Since then, this Australian lady-
Phenacoccus manihoti Matile-Ferrero in Africa bird beetle has been commonly used in various
(Neuenschwander 2001), the mango mealybug, countries on diverse crops (Copland et al. 1985;
Rastrococcus invadens (Williams) in West Africa Franco et al. 2009), and is actually one of the few
220 M. Mani and C. Shivaraju
species of natural enemies commercially avail- of the migration of natural enemies from a long
able for the biological control of mealybugs by distance. Therefore, information about the natural
means of augmentative tactics. Augmentative enemy density, late in the season, may be achieved
releases of L. dactylopii and C. montrouzieri by setting up traps baited with mealybug colonies,
against Pl. citri have been reported to be effective with or without the sex pheromone (with respect
in several Mediterranean countries and in other to each individual case).
citrus-growing areas, such as Australia and Application of chemicals alone does not solve
California. However, Mendel et al. (1999) released the mealybug problem in many cases. Many a
5,00010,000 individuals of L. dactylopii or time, more than one control method is needed to
10,00050,000 individuals of A. pseudococci per manage the mealybugs.
hectare and obtained no signicant impact both
on the mealybug infestation and on the fruit dam-
age. When the mealybug population is low, the
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Insecticide Resistance and Its
Management in Mealybugs 17
T. Venkatesan, S.K. Jalali, S.L. Ramya,
and M. Prathibha
Mealybugs throughout the world cause a variety used only once and hence pesticide applications
of economic problems and are called hard to kill are required to be repeated many times. This situ-
pests of fruit trees. Insecticides also play a criti- ation may lead to the development of insecticide
cal role in the management of mealybugs. resistance. As a result, many insecticides had
Currently, a fairly broad selection of insecticides failed to check the mealybugs. Further, different
namely organophosphates, carbamates, neonic- classes of insecticides provide various active
otinoids, insect growth regulators and keto-enols ingredients and their efcacy may be reduced if
are being used against mealybugs. However, insecticide resistance develops.
many insecticides are ineffective due to the cryp- Many insecticides do not provide adequate
tic behaviour of mealybug, its typical waxy body control of mealybugs because of the inherent
cover and clumped spatial distribution pattern. nature of mealybugs. Mealybugs are noted for the
Waxy coatings protecting the eggs, nymphs and production of dermal wax secretions. Adult
adults of mealybug make it almost impossible for mealybugs and the nymphal instars are covered
insecticides to reach them, and also there is a pos- with a waxy coating. In addition, the eggs of
sibility of development of insecticide resistance. mealybugs, protected by the waxy lamentous
Furthermore, available pesticides in the market secretions of the ovisac, are almost impossible to
may not be adequate to manage the mealybugs if reach with insecticides.
Eggs protected with waxy ovisac Adult mealybug covered with Mealybug crawlers not
waxy coating covered with waxy coating
The waxy secretion is the most common con- cals are used to control the mealybugs. However,
spicuous trait of the mealybug family. It is a crawler stage is not covered with wax, and hence,
complex system that serves different functions. this is perhaps one of the most susceptible stages
It is produced by the epidermal wax glands and of mealybug to chemicals. However, the crawler
transported to the body surface via ducts, pores stage is available only for a few days. Control
and secretory setae of various types (Foldi 1983; failure does not always imply resistance, and it is
Gullan and Kosztarab 1997). The main compo- wrong to conclude that the mealybugs have
nents of the wax of ve mealybug species developed resistance to insecticides. Mealybugs
(Planococcus citri (Risso), Pl. cus (Signoret), most frequently cause concern by their presence
Pl. vovae, Pseudococcus cryptus (Hempel) and on horticultural crops destined for export mar-
Nipaecoccus viridis (Newstead) were trialkyl kets. However, relatively few insecticides can be
glycerols and wax esters. The wax cover is used on fresh fruits without exceeding residue
believed to prevent water loss. The hydrophobic tolerances set by those markets. This restricted
property of the wax enables the mealybugs to range of permitted insecticides increases the risk
escape drowning or becoming swamped by of resistance and the potential economic impact
water in their typical cryptic sites. They are cov- that it could bring about. Flaherty et al.. (1982)
ered with a powdery wax that repels water-based reported that insufcient control measures and
insecticide solutions. The ovisac, which is also a multivoltine nature of mealybugs may speed up
wax secretion, is considered to be an adaptation the development of insecticide resistance. If an
that protects the offspring from both wet and dry insecticide has provided good control sometime
conditions, and it may also provide an attach- back but fails to effect adequate control of the
ment to the host plant. Tubular ducts and multi- mealybugs over the use of the chemical for a
locular disc pores, respectively, produce long number of years, then it is concluded that the
hollow and shorter curled laments, which make mealybug has developed resistance to that par-
up the ovisac and the male cocoon (Cox and ticular insecticide. Many a time it is not so in the
Pearce 1983; Foldi 1983). The white wax of case of mealybugs. Mealybugs are capable of
mealybugs is strongly light reective and may becoming resistant to insecticides, and it should
reduce desiccation. In some cases, the wax also not be assumed resistant until tenfold of resis-
serves to cover the honeydew droplets and to tance is observed (Valles et al. 1997; Khan et al.
protect the mealybugs from contamination by 2013). However, documentation of insecticide
their own honeydew and defensive exudates resistance in mealybugs across the globe is very
(Gullan and Kosztarab 1997). Normally, chemi- scanty.
17 Insecticide Resistance and Its Management in Mealybugs 225
17.1 Monitoring of Insecticide et al. 2005; Khan et al. 2013). Further, knowledge
Resistance of the target pest populations with respect to their
susceptibility levels to different insecticides and
While insecticides have greatly improved human how much they vary among locations can be
health and agricultural production worldwide, taken as a clue to the genetic potential for resis-
the use of insecticides has been limited by the tance development. Hence, there is a need to
evolution of resistance in many major pests, document the geographical variation of natural
including few that became pests as a result of the populations of mealybug susceptibility to insecti-
application of insecticides (Mallet 1989). cides. Further, determining a diagnostic dose for
An important component of resistance man- each test insecticide from the generated baseline
agement strategies is the ability to effectively data is needed for facilitating future resistance
monitor susceptibility levels in pest populations monitoring of mealybug (Sanderson and Roush
to insecticides. Determining dosage response of a 1992; Denholm et al. 1996).
target pest populations or species to particular
insecticides would be very useful for monitoring
insecticide resistance in mealybugs. In general, 17.2 Dosage Mortality Test
various metabolic resistance mechanisms work in Mealybugs
by detoxifying insecticides through oxidative or
hydrolytic reactions to reduce or eliminate the For establishing baseline data to different insecti-
toxic activity of insecticide. This is a dynamic cides, different stages of mealybugs were treated
process in insect populations where resistance as per the mode of action of each insecticide.
levels rise and fall according to exposure regimes Mixed stages including immature and adults of
and selection pressures (Castle et al. 1996; the mealybugs were tested for susceptibility to
Horowitz et al. 2002), which ultimately increases chlorpyrifos, dimethoate, methomyl, and imida-
the resistant individuals in the eld. Georghiou cloprid. Only immature stages were tested with
and Taylor (1986) suggested that a number of buprofezin because of its activity as a chitin syn-
factors inuence resistance development in pest thesis inhibitor that interferes with the develop-
populations including biological, ecological, ment of immature stages of susceptible insects
genetic and operational. Regular exposure to pes- (Nilima et al. 2012). There are different bioassay
ticides allows selection of individuals that are techniques which are followed for determining
naturally resistant to pesticide and develop resis- the baseline toxicity of different insecticides.
tance to survive. When a pesticide is used for the
rst time, a small proportion of the pest popula-
tion may survive exposure to the material due to 17.3 Petri Dish Bioassay
their distinct genetic makeup. These individuals
pass the genes that are responsible for resistance Susceptibility to contact insecticides (e.g. bupro-
to the next generation. Subsequent uses of the fezin, chlorpyrifos, dimethoate and methomyl)
pesticide increase the proportion of less suscep- that are applied on foliar was assessed using an
tible individuals in the population. Through this established petri dish technique (Prabhaker et al.
process of selection, the population gradually 2006a, b; Nilima et al. 2012). Morishita (2006)
develops resistance to the pesticide. also followed plastic petri dish technique in
Lack of systemic management plans may also which rst-instar nymphs of Planococcus
account for the development of resistance kraunhiae (Kuwana) were transferred onto a kid-
(Bushra et al. 2014). Different agro-ecological ney bean leaf on 1.5 % agar gel in a plastic petri
factors such as the presence of refugia, which dish. A petri dish which contained 2040 nymphs
harbour less resistant or susceptible individuals, was sprayed once with 6 ml of insecticide through
could dilute resistant gene frequencies (Sayyed a spraying tower on day 2 (rst instar), day 9
226 T. Venkatesan et al.
(second instar), day 16 (third instar) or day 25 ple, rates as low as 48 g/ha (active ingredient,
(adult), and examined for susceptibility to a.i.) of ethyl parathion provided grape mealybug
insecticide. control (Frick 1952). Eventually, these materials
became less effective. Ps. maritimus developed
resistance to parathion in the San Joaquin Valley,
17.4 Systemic Bioassay Technique California, USA (Flaherty et al. 1982). A similar
parathion resistance in the mealybugs had been
To determine baseline toxicity data of imidaclo- reported in South Africa (Myburgh and Siebert
prid, a systemic uptake technique was used as 1964). It has been reported that there could be a
described by Prabhaker et al. (2006a) and Nilima potential for resistance to buprofezin and hence it
et al. (2012). Discriminating doses are expected is recommended for twice in a year around the
to kill 100 % of a susceptible population but 0.1 world.
% of resistant individuals (French-Constant and
Roush 1990).
17.5.3 Planococcus citri (Risso)
from susceptibility tests (Misumi et al. 1994). against mealybugs in Tunisian vineyards. Mixed
Planococcus kraunhiae were collected from life stages of Pl. cus were tested for susceptibil-
places with conventional insecticide spraying and ity to all insecticides except for buprofezin,
an insecticide-free orchard. LC50 of the rst- which was measured against early and late instars
instar Hashimoto population and the resistance (rst, second and third). Variations in susceptibil-
ratio at LC50 of the other populations collected ity to each insecticide among sample sites showed
from conventional spraying orchards to that of a 7-fold difference for buprofezin, 11-fold to
the insecticide-free orchard were, respectively, chlorpyrifos, 9-fold to dimethoate, 24-fold to
0.637 ppm and 8.012.2 for cypermethrin, 1.15 methomyl and 8.5-fold to imidacloprid (Nilima
ppm and 6.07.8 for methidathion and 0.029 ppm et al. 2012).
and 15.420.2 for acetamiprid. The susceptibility
to prothiofos and methidathion decreased as the
growth stage advanced, whereas susceptibility to 17.5.7 Planococcus minor
acetamiprid remained high (Morishita 2006).
Thirumurugan and Gautam (2001) determined
the LC50 values for different insecticides and the
17.5.5 Maconellicoccus hirsutus relative resistance of Planococcus minor
(Maskell) (Pl. pacicus Cox) to various insecti-
The adult pink mealybug, Maconellicoccus hir- cides in relation to predatory beetle, Scymnus
sutus (Green), showed resistance to lambda- brunnescens (Motsch). The predators were more
cyhalothrin, pirimiphos-methyl, triazophos, resistant to endosulfan (0.07 %) than mealybugs.
pronil and decamethrin when tested under
laboratory and semi-eld conditions (Anand
and Ayub 2000). Eggs, crawlers, early nymphs, 17.5.8 Phenococcus solenopsis
late nymphs and adults of the pink hibiscus
mealybug, M. hirsutus, were tested for their In Pakistan, Bushra et al. (2014) reported insecti-
susceptibility to methyl bromide in 2-h labora- cide resistance in P. solenopsis (Tinsley) for
tory fumigations at ambient conditions (25 C, selected organophosphates and pyrethroids. The
95 % RH). Based on probit analysis of dose resistance ratio were in the range of 2.713.3-
response data, no signicant differences were fold for chlorpyrifos, 11.630.2-fold for profeno-
observed among susceptibilities of the crawler, fos, 10.646.4-fold for bifenthrin, 5.825.2-fold
early-stage or late-stage nymphs or adults at for deltamethrin and 4.125.0-fold for
either the LC50 or LC99 level, but late-stage lambda-cyhalothrin.
nymphs were more tolerant than early-stage
nymphs in a separate paired comparison test
(Zettler et al. 2002). 17.6 Resistance Management
and Prevention Strategy
reduce mealybug numbers and improve insecti- detoxifying enzymes and molecular mechanism
cide coverage. Details are provided below: of resistance by detecting different insecticide
resistance alleles, namely sodium-gated channel,
Mark spot infestations of mealybugs during KDR (Knock down resistance) and Ache.
harvest. Conne sprays to the infested crop
area: do not spray shelter or other areas around
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Mealybug Alikes
18
M. Mani and Shaheen Gul
The insects belonging to the family Pseudococcidae also called mealy scales. Yet another group belong-
and Putoidae are called true mealybugs (Williams ing to the genus Ceroplastodes is also being
2004). There are several insects (Coccoidea) simi- wrongly quoted as mealybug (Bhatnagar et al.
lar in appearance, and they are to be called as 1984). The scale insect Stictococcus vayssierei is
mealybug alikes only. By mistake, many of the also commonly called as mealybug-infested cas-
scales belonging to genera Drosicha, Icerya, sava. The following scales are also mistaken as
Perissopneumon (Margarodidae) and Pulvinaria, mealybugs and come under the category of mealy-
Chloropulvinaria, Megapulvinaria, Ceroplastodes bugs look-alike. Icerya genistae (Hempel),
(Coccidae), Dactylopius (Dactylopiidae), Pulvinaria acericola (Walsh and Riley), Pulvinaria
Eriococcus (Eriococcidae), Stictococcus ericicola (McConnell), Pulvinaria psidii
(Stictococcidae), etc. were quoted as mealybugs in (Maskell), Pulvinaria urbicola (Cockerell),
literature. In India, Drosicha mangiferae (Green) Neopulvinaria innumerabilis (Rathvon),
belongs to the Margarodidae and is popularly Philephedra tuberculosa (Nakahara & Gill),
called as mango mealybug, but truly speaking, it Protopulvinaria pyriformis (Cockerell),
should not be called as mealybug. Many species Milviscutulus mangiferae (Green), Ceroplastes
belonging to the genus Icerya (Margarodidae) are ceriferus (Fabricius), Ceroplastes rusci (Linnaeus),
also called mealybugs, like the cottony Icerya Ceroplastes cirripediformis (Comstock),
aegyptiaca (Douglas), and is wrongly called as Ceroplastes dugesii (Lichtenstein), Ceroplastodes
breadfruit mealybug in Pacic Atolls. Another cajani (Maskell), Ceroplastes oridensis
group belonging to genera Pulvinaria, (Comstock), Ceroplastes rubens (Maskell),
Chloropulvinaria, Megapulvinaria, etc. are also Eriococcus azalea (Comstock), Eriococcus quer-
similar to mealybugs when they produce ovisacs, cus (Comstock) and Dactylopius confuses
and are often mistaken as mealybugs. They are (Cockerell). Gregoporia distincta sp.n.
(Eriococcidae) is wrongly named as a mealybug
from material found on a grass of the cereal type in
a reserve in the Western Caucasus (Dantsig 1979).
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
S. Gul
Sher-E-Kashmir University of Agricultural Sciences
and Technology of Kashmir, Srinagar, India
e-mail: Shaheen.gul@rediffmail.com
18.1 Breadfruit Mealybug: Icerya et al. (2006) reported Icerya aegyptiaca as mealy-
aegyptiaca bug infestation on Santalum album.
cies affected by this insect include Acacia Incidence of Drosicha mangiferae on ash-
nilotica, A. tortilis, Casuarina equisetifolia, wagandha was reported as mealybug infesta-
Dodonaea viscosa, Grevillea robusta, Morus tion in Jammu and Kashmir, India. The pinkish
alba and Mangifera indica (Sundararaj and nymphs and female adults suck the sap from
Muthukrishnan 2008). the twigs, leaf stock and also along the midrib,
and the infestation was mainly concentrated
on the terminal part of the shoot. Sumnius ves-
18.3 Perissopneumon ferox tita and Cryptochaetum were known to attack
D. mangiferae. In western Uttar Pradesh,
Perissopneumon ferox Newstead is also similar to Drosicha mangiferae has one generation a
mealybug in appearance. Rodolia fumida and year and diapauses in the egg stage in soil for
Leptus sp. were recorded from Malihabad (Singh about 7 months.
1993). Perissopneumon ferox was reported as a The so-called mango mealybug D. stebbingi
new mealybug, Perissopneumon ferox was predated by several coccinellids, but none of
(Margarodidae, Homoptera), on mangoes from these natural enemies were found to give ade-
Uttar Pradesh, India. Heavy infestation of the quate control of D. stebbingi (Rahman and Latiff
pseudococcid P. ferox on mango was seen in 1944, Wadi and Batra 1964, Singh 1993).
Uttar Pradesh, India. Two predators, the cocci- Beauveria bassiana was found infecting nymphs
nellid Rodolia fumida and the Erythraeid Leptus of Drosicha mangiferae in the eld, and the
sp., were seen preying on P. ferox (Srivastava and pathogen was found infecting the margarodid in
Verghese 1985). Perissopneumon tamarindus orchards in ve localities in India. In eld trials
Green was also reported as a mealybug on ber on infested mango panicles, spray application of
and other crops in India (Butani 1973). a suspension having 4.8 106 conidia/ml reduced
populations of D. mangiferae by 33.3100 % in
10 days (Srivastava and Fasih 1988).
18.4 Drosicha spp. An integrated approach involving cultural,
mechanical and chemical methods is ideal for the
Drosicha stebbingi Green and Drosicha mangif- management of mango mealybugs. Raking of
erae are popularly called mango mealybugs soil four times (May, June, August and October)
besides Perissopneumon ferox. Mealybug alikes, in Uttar Pradesh afforded the best control of egg
Drosicha mangiferae (Green) and Drosicha dal- hatching of the margarodid Drosicha mangiferae;
bergiae Green was recorded as mealybugs on 30 % of the eggs hatched, as compared with 68 %
pomegranate and papaya in India. Mealybug for no treatment (Chandra et al. 1989). Complete
alikes, Drosicha stebbingi Green, D. mangiferae control could be obtained by the use of grease
(Green) (Pruthi and Batra 1960), Drosichiella bands round the trunks from the second week of
tamarindus Green and Perissopneumon tama- December. An alternative method proved to be
rindus Green (Butani 1973) were reported as banding with coal tar, which remains effective for
mealybugs on ber in India. Drosicha stebbingii only a relatively short period (Prasad and Singh
Green on forest plants is reported as a mealy 1976). Sticky bands were found to remain effec-
bug occurring throughout the sal (Shorea tive for only a short time (up to 15 days after
robusta) forests of north India. It was also application). The commonly used bands of mix-
quoted as mealybug on Tectona grandis and ture of rosin and castor oil (4:5) and coal tar and
Albizia spp. in India (Joshi 1992). Drosicha grease (2:1) prevented the nymphs from ascend-
mangifera was also recorded as a mealybug pest ing for only up to 56 days after application.
on black nightshade (Solanum nigrum L) and Field tests were carried out in Hissar, India, and
Indian gooseberry (Phyllanthus emblica) from revealed that the slippery band of alkathene sheet
Uttar Pradesh, India. was most effective of all in blocking the ascend-
234 M. Mani and S. Gul
ing nymphs, as an average of 2.79 nymphs per effective against gravid females of the pest.
sample area were able to cross it every alternate Spraying of acephate, methyl demeton, mono-
day as compared with 407.3 nymphs on untreated crotophos, quinalphos, dimethoate and phos-
trees (Lakra et al. 1979). A 30-cm-wide polyeth- phamidon at 0.08 % was able to keep the
ylene band tied round the tree 50100 cm above population of mealybug D. mangiferae under
the ground and with its lower edge plastered over check.
with mud was sufciently slippery to prevent the
passage of Drosicha stebbingi nymphs and much
cheaper than the conventional sticky band (Bindra 18.5 Stictococcus vayssierei
and Sohi 1974). The double girdle band of alk-
athene was more effective as it stopped the few Stictococcus vayssierei Richard has been reported
nymphs of Drosicha mangiferae that managed to as root mealybug of cassava (Manihot esculenta)
cross the rst band (Srivastava 1980). in Cameroon (Ngeve 2003). The larvae and adults
Trunk sprays of quinalphos, diazinon and attack young feeder roots of germinating cut-
methyl parathion at 0.0.750.15, 0.050.1 and tings, causing extensive leaf-fall, wilting, tip die-
0.050.1 % in Haryana were highly effective back and death of plants. Plants that escape early
against ascending rst instar nymphs that had infestation develop normally and tuberize, but the
collected below bands. In Bihar, alkathane mature tuberous roots are small and become cov-
banding was followed by three to four applica- ered with the root scale, making them unattract-
tions to the trunk of 0.04 % malathion or three ive to market. In severe infestations, a mature
of 0.03 % dimethoate, 0.03 % phosphamidon, tuberous root of about 40 cm long may harbour
0.04 % diazinon or 0.05 % thiometon during up to 500 mealybugs. It is most severe during the
January. All these insecticides were equally dry season in lateritic and clayey soils, in elds of
effective when applied to the shoots (after band- depleting fertility and in thinly prepared land
ing of the trunks) in late February and early where planting has been done on the at. The
March. Diazinon and thiometon were too expen- prevalence of the pest in the semi-humid forest
sive for their use to be recommended (Prasad region of Cameroon increased from 12.5 % in
and Singh 1976). Field tests carried out in Delhi 1990 to 87.5 % in 1999. S. vayssierei infestation
indicated that diazinon was the most effective was more severe (30 mealybugs/hill) when cas-
compound and was signicantly superior to sava was planted on the at than when planted on
monocrotophos and chlorpyrifos, both of which, ridges (16 adults/hill). Plants also sprouted better
however, gave fairly satisfactory control of the (91 %) when cassava was planted on ridges than
pest (Srivastava 1980). Infested trees were when planted on the at (71 %). Root yields (31.4
sprayed once, with acetamiprid at 100 g/100 L t/ha) and root numbers (7 roots/hill) were also
of water against rst instar in the second week higher in cassava planted on ridges than in those
of February in Pakistan (Karar et al. 2009). grown on the at (24.5 t/ha and 4.5 roots/hill,
Against Drosicha mangiferae on guava, fenitro- respectively). For plants grown on the at, the
thion at 0.1 % was the most effective treatment, improved clones suffered the least attack by S.
followed by phosalone at 0.07 %, quinalphos at vayssierei, clones 8017 and 8034 showing the
0.05 %, monocrotophos at 0.04 %, parathion- most tolerance (19 and 22 females/hill, respec-
methyl at 0.05 %, and bromophos-ethyl at 0.07 tively) when compared with the local, Meyiboto
% and phosphamidon at 0.03 %. Phenthoate at (49 females/hill). Stictococcus vayssierei was
0.05 %, dimethoate at 0.03 % and malathion at more severe when cassava was intercropped;
0.1 % were less effective (Dalaya et al. 1983). there were 40, 48 and 59 mealybug adults per hill
Among 24 insecticides that were tested against when cassava was intercropped, respectively,
Drosicha mangiferae in Haryana, quinalphos at with maize, groundnuts or maize and groundnuts
0.025 % and fenitrothion, carbophenothion and combined. By contrast, maize suffered no yield
parathion-methyl, each at 0.05 %, were highly depression when intercropped with cassava. S.
18 Mealybug Alikes 235
and foliar spray, respectively, plays a signi- Prasad VG, Singh RK (1976) Prevalence and control of
mango mealy bug Drosicha stebbingi (Green) in
cant role in suppression of the pest (Shaheen
Bihar. Indian J Entomol 38(3):214224
et al. 2014). Pruthi HS, Batra HN (1960) Some important fruit pests of
North West India, Indian Coucil Agricultural Rearch
(ICAR) Bull No. 80, New Delhi 113 p
Rahman KA, Latiff AM (1944) Description, bionomics and
control of the giant mealybug Drosicha stebbingi
References (Homoptera, Coccidae). Bull Entomol Res 35:197209
Rajabalee A, Banymadhub N (1990) Preliminary notes on
Bhatnagar VS, Jadhav DP, Pawar CS (1984) Parasitoids of the mealybug Pseudaspidoproctus fulleri Cockerell
pigeonpea mealy bug, Ceroplastades cajani Mask. Int (Homoptera: Margarodidae), a new introduction to the
Pigeonpea Newslett 3:45 Island of Mauritius [French]. Revue Agricole et
Bindra OS, Sohi BS (1974) A note on control of the Sucriere de lIle Maurice 69(13):6061
mango mealybug in the Punjab. Indian J Hortic Rao VP (1950) Iceryine scale insects recorded from the
31:102103 orient. Indian J Entomol 12:3966
Butani DK (1973) Insect pests of fruit crops and their con- Shaheen G, Zahoor B, Ahmad AS (2014) Management of
trol. Pesticides 7:3335 almond mealy bug, Drosicha dalbergiae Stebbing by
Chandra A, Bhati DPS, Singh KM (1989) Note on the chemical intervention. Ann Plant Protect Sci 22(1):2226
effect of soil raking and irrigation on survival and Singh SP (1993) Biological control of pests. In: Chadha KL,
hatching of eggs of mango mealy bug, Drosicha man- Pareek OP (eds) Advances in horticulture III. Malhotra
giferae Green. Curr Agric 13(12):103104 Publishing House, New Delhi, pp 15911616
Dalaya VP, Rajput SG, Mali AR, Mohite PB (1983) Srivastava RP (1980) Efcacy of alkathene bands to pre-
Comparative efcacy of insecticides against guava vent ascent of mango mealybug nymphs on mango
mealy bug Green. Indian J Plant Protect trees. Indian J Entomol 42(1):122129
11(1/2):138139 Srivastava RP, Fasih M (1988) Natural occurrence of
Dantsig EM (1979) A new peculiar genus of mealybugs Beauveria bassiana, an entomogenous fungus on
(Homoptera, Coccoidea, Eriococcidae) from the mango mealybug, Drosicha mangiferae Green. Indian
European part of the USSR [Russian]. Trudy J Plant Pathol 6(1):810
Vsesoyuznogo Entomologicheskogo Obshchestva Srivastava RP, Verghese A (1985) Record of a new mealy-
61:4647 bug, Perissopneumon ferox Newstead (Margarodidae:
Joshi KC (1992) Handbook of forest, zoology & entomol- Homoptera) on mango from Uttar Pradesh, India.
ogy. Oriental Enterprise, Dehradun, 383p Entomon 10(2):184
Karar H, Arif MJ, Sayyed HA, Saeed S, Abbas G, Arshad Sundararaj R, Karibasavara LR, Sharma G, Muthukrishnan
M (2009) Integrated pest management of mango R (2006) Scales and mealybugs (Coccoidea:
mealybug (Drosicha mangiferae) in mango orchards. Hemiptera) infesting Sandal (Santalum album Linn.).
Int J Agric Biol 11:8184 Entomon 31(3):239241
Lakra RK, Kharub WS, Singh Z, Lakra RK, Kharub WS, Sundararaj R, Muthukrishnan R (2008) Sucking pest
Singh Z (1979) Comparative efcacy of some banding complexes of sandal and their host range in South
materials against mango mealybug, Drosicha mangif- India. In: Gairola S, Rathore TS, Joshi G, Arun Kumar
erae Green. in Haryana. Indian J Entomol AN, Aggarwal P (eds) Proceedings of the national
41(2):170176 seminar on conservation, improvement, cultivation
Malik RA, Punjabi AA, Bhat AA (1972) Survey study of and management of sandal (Santalum album L.).
insect and non insect pests in Kashmir. Horticulture Brilliant Printers, Bangalore, pp 111120
3:2944 Wadi SR, Batra HN (1964) Pests of tropical and subtropi-
Masoodi MA, Bhagat KC, Koul VK (1988) Drosicha dal- cal fruit trees. Entomological society of India, New
bergiae Green (Coccidae: Homoptera) a new pest of Delhi, pp 464466
almond in India. Ann Biol 4:12 Waterhouse DF (1991) Possibilities for the biological
Ngeve JM (2003) The cassava root mealybug (Stictococcus control of the breadfruit mealybug, Icerya aegyptiaca,
vayssierei Richard) (Homoptera: Stictococcidae): a on Pacic Atolls. Micronesica (Suppl. 3):117122
threat to cassava production and utilization in Williams DJ (2004) Mealybugs of southern Asia. The
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London/Kuala Lumpur, 896 p
Part II
Management of Mealybugs in Agricultural
and Horticultural Crops
Rice
19
Gururaj Katti
19.1 Mealybug Species rice mealybug species Brevennia rehi has under-
gone several modications across space and time
Mealybugs are injurious to rice in several coun- (CABI 2003). At rst, the rice mealybug
tries. Among the species, Brevennia rehi Brevennia rehi was recorded as Ripersia sacchari
(Lindinger) is widely distributed across the world Green by Lefroy (1908), attacking rice in India,
in South and South East Asia, North America and and later, Brevennia rehi was conrmed as the
Australasia (Table 19.1). In a recent report also, valid name for the rice mealybug by Miller (1975).
the rice mealybug has been listed as one of the It has become a primary pest in Bihar, and also in
important pests of rice in Bangladesh (Ahmad other rice-growing states such as West Bengal,
et al. 2011). It is found to cause heavy loss to the Orissa, Bihar, Andhra Pradesh, Tamil Nadu,
growers in India and Pakistan. Identication of Karnataka, Kerala and Maharashtra (CIE 1979).
Table 19.1 List of mealybug species recorded on rice in different regions of the world
Species Country/Region Reference
Brevennia rehi (Lindinger) Israel, Iraq, Azerbaijan, Tajikistan Ben-Dov (2008)
and Brazil
Syns: Heterococcus rehi (Lindinger); Bangladesh Alam et al.(1979), Alam and
Heterococcus tuttlei Miller and Bhuiyan (1964)
McKenzie; Rhizoecus cynodontis Nepal Pradhan (1981)
Bodenheimer; Brevennia femoralis
Taiwan Liu and Tao (1988)
Borchsenius
Australia and Papua New Guinea Williams et al. (1981)
India Narayan and Ram (1985), Radja
(1985), Velusamy and Babu
(1986), Gopalan et al. (1987a),
Ghode and Mishra(1988),
Lakshmanan et al. (1988),
Raguraman et al. (1991), Jayarani
and Velusamy (1994)
Bangladesh, Pakistan and Williams (2004)
Philippines
USA Miller and McKenzie (1970)
Chlorozococcus mireorum Matile Cameroon Ben-Dov (1994)
Ferren
Chorizococcus ilu Williams Williams (1970)
Bangladesh Alam and Karim (1981)
Dysmicoccus boninsis (Kuwana) Taiwan and China Liu and Tao (1988)
USA Miller and McKenzie (1970)
Bangladesh Alam and Karim (1981)
Dysmicoccus brevipes (Cockerell) Williams (1970)
Dysmicoccus oryzae (Wijati) Java Williams (2004)
Bangladesh Alam and Karim (1981)
Formicoccus lingnani (Ferris) Malaysia and Thailand Williams (2004)
Geococcus oryzae (Kuwana) Williams (1970)
Bangladesh Alam and Karim (1981)
Nipaecoccus graminis (Maskell) Williams (1970)
Bangladesh Alam and Karim (1981)
Novaniliacoccus oryzae Ghosh & India Williams (2004)
Ghise
Paracoccus ilu (Williams) Fiji and New Zealand Ben-Dov (2008)
Planococcoides lingnani (Ferris) China Ben-Dov (2008)
Bangladesh Alam and Karim (1981)
Planococcus minor (Maskell) Philippines Williams (2004)
Pseudococcus saccharicola Takahashi Australian and Oriental region Ben-Dov (2008)
Bangladesh Pathak and Khan (1994)
Malaysia and Thailand Williams (2004)
Pseudorhodania oryzae Tang China Ben-Dov (2008)
Saccaharicoccus sacchari (Cockrell) Williams (1970)
Bangladesh Alam and Karim (1981)
Trionymus ceres Williams India and Pakistan Williams (1970; 2004)
Bangladesh Alam and Karim (1981)
19 Rice 241
stages of the rice crop. The infested plants appear develop properly and have a bitter taste; if present
stunted and scorched. High incidence inhibits in normal food, they spoil the avour after being
panicle emergence, and plants may even dry. cooked. The pest is also known to transmit the
Grains from mealybug-infested plants do not virus known as chlorotic streak (Williams 2004).
19.3 Factors Inuencing the rice nurseries, which provide the main source
Incidence of Mealybugs of infestation. The alternative hosts include
Cynodon dactylon, Cyperus rotundus,
The prevalence of dry period, presence of grassy Echinochloa crus-galli, Echinochloa colona,
weeds, well-drained soils and upland/rain-fed Panicum repens and Paspalum scrobiculatum.
environments are major factors inuencing the The mealybug damage is found mainly conned
mealybug incidence. Increased temperature and to upland and rain-fed environments, particularly
wind velocity and decreased relative humidity in elds with uneven soil surface where the plants
have been reported to increase the incidence of B. grow in relatively dry soil patches. It occurs in
rehi (Radja 1985). Also, the pest infestation is great number during the rainy season.
more severe in unirrigated and upland elds
(Mammen 1976; Pradhan 1981). The planting
dates and irrigation regimes also inuenced the 19.4 Extent of Losses
incidence of rice mealybugs. Early planting and
continuous pounding of irrigation water at 5-cm The rice mealybugs cause heavy losses to crops
depth throughout the growing period resulted in in Bangladesh, India, and Thailand. High density
lower intensity of rice mealybug infestation (>100 mealybugs/hill) causes plants to wilt and
(Gopalan et al. 1987a). Also, type and dosage of die. Despite being a traditional pest in the upland
nitrogenous fertiliser applied affected infestation paddy in the eastern states of Orissa and West
levels: higher levels of nitrogen increased the rice Bengal, there are few reports on the quantica-
mealybug infestation, whereas the application of tion of the extent of rice mealybug incidence or
raw coir pith, raw sugarcane trash and farmyard its damage in India. Banerjee (1956) reported
manure reduced the infestation (Backialakshmi incidence of mealybugs from Midnapore, Nadia,
1994). During the off season, rice mealybugs sur- 24 Parganas, Bankura, Murshidabad and
vive on a variety of grasses, later spreading into Jalpaiguri districts of West Bengal. Satpathi et al.
242 G. Katti
(2005) reported an average damage up to 7 % in timely destroying of the infested plants are useful
the rice-growing areas of West Bengal. Ghode in preventing its spread and impact.
and Mishra (1988) reported a serious outbreak of
its occurrence in Dhenkanal, Cuttack and Puri
districts of Orissa state, the affected areas being 19.6 Varietal Resistance
4,000 ha, 2,780 ha and 303 ha, respectively.
Velusamy and Babu (1986) observed a severe Traditional/local varieties and improved culti-
attack of mealybugs in an area of 100 ha of rice in vated varieties showed low levels of resistance
the Pudukottai district of Tamil Nadu, India. The (Alam et al. 1979; Heinrichs 1983). Radja (1985)
population of adults and nymphs was 650750/ and Gopalan et al. (1987b) reported varieties
hill and the affected plants failed to produce pan- such as IET 8616, AS 89090 and IET 12798 with
icles. Later in two villages of Tamil Nadu, low infestation after screening them under eld
extremely severe incidence of mealybug popula- conditions. Mallikarjuna Rao (1987) found that
tions was reported up to 91.1 per tiller (Nalini TNAU 80030, TM 1087 and CO 43 were toler-
et al. 2011). There was yield reduction in ant, with the outer leaf tip turning yellowish,
extremely stunted rice plants at a population level despite high bug population, while TNAU 831520
of 50 mealybugs/hill (Backialakshmi 1994). The and TNAU 831521 were found to be resistant and
association of high mealybug incidence with the moderately resistant, respectively. Further stud-
occurrence of sheath rot disease further aggra- ies identied more resistant sources such as Ptb
vates the yield reducing potential of this pest in 33, IR 56 and IR 58, Tending, Badal 2, Rathu
rice (Alam and Karim 1981; Lakshmanan et al. Heenati, Ptb 21, Sufaida 172, IR 42 and IR 72,
1988; 1991). Rice mealybugs are also associated Senawee, Sufaida 172, DR 52 and ARC 575
with rice chlorotic streak viruses as the transmis- (Jayarani 1992: Jayarani and Velusamy 1994:
sion studies with the bug were positive. There Backialakshmi 1994). The studies on resistance
have been reports of widespread and severe out- mechanism indicated that feeding by rice mealy-
breaks of rice mealybug infestation with associa- bug resulted in a marginal increase in total phe-
tion of sheath blight and sheath rot diseases in nolic content and a large increase in total sugars,
Bangladesh during the drought years of 1950, reducing sugars, non-reducing sugars, isoleucine
1957, 1966, 1972 and 1979 (Alam et al. 1979). and proline content of the rice plants (Gopalan
Both traditional and improved varieties showed et al. 1987c). Resistant varieties had low total
infestation, and the crop losses were estimated at nitrogen, low potassium and high calcium con-
30 % because of the combined effects of drought tents compared with the moderately resistant and
and mealybug. Pradhan (1981) mentioned B. rehi susceptible varieties (Mallikarjuna Rao 1987).
as a pest of rice in the Terai belt of Nepal which Antixenosis and antibiosis were also reported,
included areas of Sarlahi, Bara, Parsa, Rautahat resulting in low oviposition and egg hatchability,
and Dhanusha. Rice mealybug has also been slow nymphal development, reduced adult lon-
reported as tuttle mealybug-infesting Bermuda gevity and low fecundity. Steam distillate extracts
grass (Cynodon dactylon) in USA (Ben-Dov of resistant varieties adversely affected the ovipo-
2012). sitional behaviour and were toxic to crawlers
(Lakshmanan and Velusamy 1991).
19.5 Management
19.7 Cultural Control
It is difcult to control B. rehi because of its pro-
tective waxy covering over its entire body and a Removal of alternative hosts in the vicinity of the
secure position in between the stalks and leaf eld is recommended to prevent pest multiplica-
sheath; however, early detection of the infesta- tion (Ayyar 1939). It is also advised to infested
tion in the nursery as well as pulling out and plants at the post-panicle initiation stage, burying
19 Rice 243
them in the soil and replanting to prevent further parathion-methyl and demeton (Anantanarayanan
spread of the pest (Alam et al. 1979). Early plant- and Abraham 1957), malathion (Wahed 1959;
ing and regular irrigation had also resulted in Alam 1965), demeton-S-methyl (Alam 1965;
lower levels of mealybug infestation (Pradhan Mallikarjunaa Rao 1987), diazinon, phosphami-
1981; Radja 1985; Gopalan et al. 1987a). The don, fenthion and fenitrothion (Alam 1965;
application of organic products such as raw sug- Alam et al. 1979; Radja 1985; Lakshmanan et al.
arcane trash and farmyard manure reduced the 1991), dicrotophos (Alam et al. 1979), dimetho-
infestation (Backialakshmi 1994). ate (Radja 1985; Gopalan et al. 1987d; Radja
1985), monocrotophos (Mallikarjunaa Rao
1987), Chlorpyriphos and isofenphos as seed
19.8 Biological Control treatment (Rajamani et al. 1987) and phorate as
furrow treatment (Rajamani et al. 1987) were
Few natural enemies have been recorded as poten- recommended for the control of B. rehi. The
tial enemies of different stages of rice mealybugs, organocarbamate insecticides found effective
and no eld release studies have been made. The included fenobucarb (Radja 1985), carbofuran
parasitoids recorded so far are Ceraphron sp., and carbosulfan as seed treatments (Rajamani
Adelencyrtus sp., Cheiloneurus sp., Doliphoceras et al. 1987) and carbaryl as an ovicide (Gopalan
sp., Mayeridia sp., Parasyrphophagus sp., et al. 1987d).
Xanthoencyrtus sp., Rhopus fullawayi, Gyranusa
sp., Aprostocetus sp., Chrysochoris sp., Desostenus
sp., Tetrastichus sp., Lymaemon sp., Callitula sp., References
Diparini sp. and Thysanus sp., while predator spe-
cies include Anatrichus pygmaeus Lamb, Ahmad H, Ahmed A, Kamal NQ, Mia TMA (2011) Final
Report on Consulting Service for Conducting Pest
Domomyza perspicax (Knab), Leucopis luteicor-
Risk Analysis (PRA) of Rice (both Production and
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Das 1971; David and Ananthakrishnan 2004; Agriculture quarantine services strengthening pro-
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Ben-Dov Y (2008) The rice mealybug, Brevennia rehi Depart Agric Mem Ent Ser 2:111137
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Sundaram CV (1935) The soorai disease of paddy. Kerala. Int Rice Res Newslett 1(2):1415
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CIE (1979) Distribution maps of plant pests, No. 401. predators of rice mealybug, Ripersia oryzae Green
CAB International, Wallingford (Hemiptera: Pseudococcidae). Curr Sci 37:354355
David BV, Ananthakrishnan TN (2004) General and Miller DR (1975) A revision of the genus Heterococcus
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New Delhi, 1184 p Cooidea: Pseudococcidae). Tech Bull US Dept Agric
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Orissa. Plant Prot Bull (Faridabad) 40(1):35 genus Heterococcus (Homoptera: Coccoidea:
Gopalan M, Radja NC, Balasubramanian G (1987a) Pseudococcidae) with the description of a new
Effect of different planting dates and irrigation regimes species. Ann Entomol Soc Am 63:438453
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(Lindinger). Madras Agric J 74(45):225227 dence of rice mealybug Brevennia rehi (Lindinger)
Gopalan M, Radja NC, Balasubramanian G (1987b) (Hemiptera: Pseudococcidae) and its gregarious para-
Screening rice varieties for resistance to mealybug. sitoid Rhopus nigroclavatus (Ashmead) (Hymenoptera:
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Biochemical changes in rice plants infested with of rice mealybug during 1982 in Bihar. Plant Prot
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Ovicidal activity of insecticides on eggs of Brevennia International Rice Research Institute, Los Banos, 89 p
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Hayat M (1970) Studies on the genera of the family plants. Int Rice Res Newslett 6(4):1112
of Signiphoridae (Hymenoptera: Chalcidioidea) Prakasa Rao PS, Das PK (1971) New records of some
recorded from India. Entomophaga 15(4):387399 natural enemies of rice mealybug, Ripersia oryzae
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the thrips, caseworm, leaffolder. rice bugs and mealy 8:111112
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Conference, 1822 April 1983 at IRRI, Manila, rice mealybug, Brevennia rehi (Lindinger)
Philippines (Pseudococcidae: Hemiptera). MSc thesis, Tamil
Jayarani S (1992) Genetic evaluation for mealybug, Nadu Agricultural University, Coimbatore
Brevennia rehi (Lindinger) (Hemiptera: Raguraman S, Saroja R, Manickavasagam S (1991) New
Pseudococcidae) resistance in rice varieties. MSc parasitoid record on the rice mealybug, Brevennia rehi
(Ag.) thesis, Tamil Nadu Agricultural University, Lindinger (Pseudococcidae: Hemiptera). Indian
Coimbatore J Entomol 53(1):170171
Jayarani S, Velusamy R (1994) Resistance to rice mealy- Rajamani S, Pasulu IC, Dani RC, Kulshreshtha JP (1987)
bug in white backed planthopper resistant rice variet- Evaluation of insecticides and plant products for the
ies. Int Rice Res Newslett 19(2):13 control of insect pests of rainfed upland rice. Indian
Lakshmanan P, Velusamy R (1991) Resistance to sheath J Plant Prot 15(1):4350
rot (ShR) of breeding lines derived from Oryza ofci- Santhanaraman T (1952) Some experiments on the con-
nalis. Int Rice Res Newslett 16(6):89 trol of the rice stem borer (Schenobius incertulas
Lakshmanan P, Manokaran T, Mohan S (1988) Role of Wlk.) and the rice mealy bug (Ripersia oryzae Green).
mealybugs on sheath rot disease manifestation in rice Plant Prot Bull 4:83100
plants. Trop Pest Manage 34(3):356357 Satpathi CR, Mukhopadhyay AK, Katti G, Pasalu IC
(2005) Integrated management of upland rice insects
19 Rice 245
in West Bengal. (National Agricultural Technology Williams DJ (1970) The mealybugs (Homoptera,
Project) NATP RRPS-22. Monograph No. 1. BCKVV, Coccoidea, Pseudococcidae) of sugarcane, rice and
Kalyani, p 46 sorghum. Bull Entomol Res 60:109188
Velusamy R, Babu PCS (1986) Occurrence of rice mealy- Williams DJ (2004) Mealybugs of southern Asia. The
bug in Tamil Nadu, India. Int Rice Res Newslett Natural History Museum/Southdene SDN. BHD,
11(4):35 London/Kuala Lumpur, 896 p
Wahed AT (1959) Pests of paddy and their control mea- Williams DJ, Radunz LAJ, Brookes HM (1981) The rice
sures with special reference to resistance. In: mealybug Brevennia rehi (Lindinger) now recorded
Proceedings of the First International Rice Breeding from Australia and Papua New Guinea (Hemiptera:
Training Centre, Dacca, February 1628, pp 113118 Coccoidea: Pseudococcidae). J Aust Entomol Soc 20:46
Wheat
20
Srinivasa Babu Kurra, Jeyakumar Ponnuraj,
and Shyam Prasad Gogineni
Fig. 21.1 Barley mealybug and mealybug damage in the barley eld
Fig. 22.1 (a) Roots infested with mealybugs, (b) (e) Mealybugs feeding on stem, (f) Eggs inside the protec-
Mealybugs feeding on leaves, (c) Mealybugs on either tive pouch (ovisac)
sides of vein, (d) Pegs and pods infested with mealybugs,
254 G. Harish and M.V. Nataraja
groundnut kernels, and they turn into black (http:// occurred during February and March months
www.daff.qld.gov.au/26_14460.htm). (Rao and Srinivasan 1987). In Gujarat, F. virgata
was observed on harvested groundnut plants in
the month of May (Anonymous 2003). In Ghana,
22.2.4 Paracoccus marginatus Paracoccus marginatus was found infesting ground-
William and Granara de nut during July to March months (Cham et al.
Willink 2011). In Sind and Punjab provinces of Pakistan,
Phenacoccus solenopsis peak population was
Paracoccus marginatus William and Granara de observed during the rst week of September on
Willink is commonly called as papaya mealybug groundnut (Abbas et al. 2010; Lysandrou et al.
and is also known to infest groundnut, and its 2012). It was also recorded during June to October
degree of infestation recorded was below 15 % in (2012) in Junagadh district of Gujarat.
Tamil Nadu, India (Selvaraju and Sakthivel
2011). It was also reported infesting groundnut
from Akraman and Nsawam regions of Ghana 22.4 Management
(Cham et al. 2011).
Managing mealybugs on groundnut crop requires
a holistic approach through proper integration of
22.2.5 Phenacoccus solenopsis several pest management tactics such as cultural,
Tinsley mechanical, physical, biological, behavioral, and
chemical measures.
Phenacoccus solenopsis Tinsley is commonly
called as solenopsis mealybug, a polyphagous
pest known to multiply on different host plants, 22.4.1 Cultural Control
including groundnut. In Australia, it was found
attacking groundnut (http://www.daff.qld.gov. Crop should be kept free from weeds and
au/26_14460.htm). In India, it was rst time alternate hosts harboring the mealybugs.
recorded infesting GG-20, a variety of groundnut Plants should be maintained in healthy condi-
during kharif, in 2012 in Junagadh district of tion and avoid water stress.
Gujarat. These mealybugs were also found asso-
ciated with pods, pegs, green succulent stems,
and branches. 22.4.2 Mechanical Control
Singh TVK, Goud TR, Azam KM (1986) Attack of mealy- cies from southern Asia. Bull Entomol Res
bug, Dysmicoccus breviceps on groundnut. Indian 86:617628
J Entomol 48(3):358 Williams DJ (2004) Mealybugs of southern Asia. The
Tanwar RK, Jeyakumar P, Monga D (2007) Mealybugs Natural History Museum/Southdene SDN. BHD,
and their management, Technical bulletin no. 19. London/Kuala Lumpur, 896 p
National Centre for Integrated Pest Management, New Williams DJ, Granara de Willink MC (1992) Mealybugs
Delhi, 16 p of Central and South America. CAB International,
Williams DJ (1985) Australian mealybugs. British London, 635 p
Museum (Natural History), London, 431 p Williams DJ, Watson GW (1988) The scale insects of the
Williams DJ (1996) A brief account of the hibiscus mealy- tropical south pacic region, part 2: the mealybugs
bug Maconellicoccus hirsutus, a pest of agriculture (Pseudococcidae). CAB International Institute of
and horticulture, with descriptions of two related spe- Entomology, Wallingford, 260 p
Sunower
23
K.S. Jagadish, Chandrashekar, H. Basappa,
G. Basana Gowda, and Y.G. Shadakshari
In recent years, the mealybugs are found to infest likelihood of severe incidence of this mealybug
sunower crop in different sunower-growing on sunower crop in Punjab, Haryana and Gujarat
regions. Phenacoccus solenopsis (Tinsley) whenever this pest causes severe damage to cot-
(Basappa 2007; Jagadish et al. 2008), ton crop (Basappa 2008).
Maconellicocus hirsutus (Green) (Basappa 2008;
Rathod et al. 2008) and Paracoccus marginatus
(Williams and Granara de Willink) (Jagadish 23.1 Bionomics
et al. 2010) are known to attack sunower in
India. In Australia also, Ph. solenopsis was found The total developmental period of egg to adult P.
attacking sunower (http://www.daff.qld.gov. solenopsis on sunower was completed in 2030
au/2614460.htm). Phenacoccus madeirensis days and the fecundity of female was about 500
(Green) is also known to infest sunower (Bend- 650 nymphs (Basappa 2008; Chandrashekar
Dov 1994). 2011; Anonymous 2011). Around Bengaluru,
Phenacoccus solenopsis is the major mealy- India, the mealybug infestation initially appeared
bug species attacking sunower particularly in the rst week of January and gradually
grown nearer to cotton in India. It is found severe increased as the season advanced. Then it
on sunower in different parts of Karnataka increased abruptly to reach 156.20 in the 15th
(Bengaluru, Chitradurga, Bellary, Haveri, standard week (915 April). Later, the mealybug
Bagalkot, Koppal and Gadag districts), Andhra population declined gradually and reached
Pradesh (Nandyal, Kadri, Anantapur, Karimnagar, 113.78 in the 18th standard week (30 April5
Gouraram, Ranga Reddy and Hyderabad), May). The mealybug population remained nil
Maharasthra (Akola, Aurangabad and Jalna dis- during kharif and rabi seasons (JuneDecember)
tricts) and Tamil Nadu (Coimbatore, Tirupur, (Chandrashekar 2011). It was signicantly posi-
Erode, Salem and Namakkal districts). There is a tively correlated with maximum temperature
(0.870**) and sunshine hours (0.509**) and neg-
atively correlated with rainfall (0.177) and
K.S. Jagadish (*) Chandrashekar G.B. Gowda
Y.G. Shadakshari morning (0.627**) and evening (0.743**) rel-
University of Agricultural Sciences, GKVK, ative humidity. The population of the encyrtid
Bengaluru 65, India parasitoid (Aenasius bambawalei Hayat) was
e-mail: jagsan_san@yahoo.co.in also signicantly positively correlated with P.
H. Basappa solenopsis population (0.985**) and weather
Directorate of Oilseeds Research, ICAR, parameters, namely maximum temperature
Rajendranagar, Hyderabad 30, India
(0.845**) and sunshine hours (0.493). The et al. 2008; Chandrashekar 2011; Basappa 2008).
regression coefcient of the mealybug and its Different ant species were found to transfer
parasitoid with weather parameters indicated that P. solenopsis from one sunower plant to another,
the population dynamics of mealybug and para- and also found to provide protection to the mealy-
sitoid was dependent on weather parameters to bug from predators, parasitoids and other natural
the extent of more than 70 % (Chandrashekar enemies (Chandrashekar 2011). Ants were
et al. 2012). responsible for quick colonization of P. solenop-
At Raichur in Karnataka, India, peak infesta- sis to newer areas (Saini et al. 2009) (Fig. 23.1).
tion of P. solenopsis was observed from October
to November, while no incidence was found from
June to September (Anonymous 2011). At 23.3 Damage
Hyderabad, India, maximum population of
mealybug was noticed on sunower from April Incidence of Ph. solenopsis is observed on sun-
to August (Basappa 2008). High temperatures ower a week after germination onwards, up to
along with low humidity are congenial for rapid maturity stage, but the level of damage varies
growth and multiplication of P. solenopsis, while with different phenological stages of the crop.
high-intensity rains and wet spells adversely Sunower is highly susceptible to mealybug
affect its infestation (Saini et al. 2009). attack in the seedling stage. One adult can cause
typical symptoms of curling of leaves, stunted
growth, deformation and death of plants, within
23.2 Ant Association 30 days of germination. At this stage, it could
with Mealybugs cause 100 % crop loss. If infestation occurs at
vegetative stage, symptoms of curling of leaf,
Ten species of ants, namely Paratrechina longi- stunted growth and deformation without death of
cornis (Latreille), Myrmicaria brunnea plant were observed, but the plant could not pro-
(Saunders), Monomorium pharaonis (Linnaeus), duce owers; however, it could cause 100 % loss
Tapinoma melanocephalum (Fabricius), in patches. If incidence is at the reproductive
Camponotus sericius (Fabricius), Solenopsis stage of the crop, it affects ower buds and ow-
geminata (Fabricius), Anoplolepis gracilipes ers, leading to deformation of head without seed
(Smith), Camponotus compressus (Fabricius), set. In some cases, partial seed setting is also
Monomorium latinode (Mayr) and Oecophylla noticed with about 50 % yield loss (Anonymous
smaragdina (Fabricius), were found attending to 2011; Basappa 2008; Chandrashekar 2011;
the mealybug P. solenopsis in sunower (Jagadish Rathod et al. 2008). The overall appearance of
Fig. 23.1 Ants attending to the mealybugs (a) Monomorium pharaonis (b) Camponotus compressus
23 Sunower 259
the plant was bushy and stunted, with the infested tors was observed from February to April
plants showing a reduction in their height, as around Bangalore (Chandrashekar 2011).
compared to the uninfested plants (Jagadish et al. Activity of natural enemies associated with
2009). Sunower yield in Tamil Nadu slipped to mealybug was more during April to June 2008,
700 kg/acre from the normal yield of 10001200 and later its population was reduced to mini-
kg/acre, mainly due to the mealybug infestation mum around Hyderabad (Basappa 2008).
in the entire growing belt (Anonymous 2009; Lecanicillium lecanii (Zimm.) Zare & Gams
Suresh et al. 2010). At Akola, Maharashtra, 60 % was found to be highly pathogenic to P. sole-
reduction in sunower seed yield was reported in nopsis. At an initial inoculum of 1 104 conidia
the case of Morden variety due to mealybug mL1, lethal time (LT50) was 3.77 and 2.51
attack by Rathod et al. (2008) (Fig. 23.2). days for nymphs and adults, respectively
(Gulsarbanu et al. 2009) (Figs. 23.3 and 23.4).
References
Anonymous (2009) Mealybug hits sunower yield. The
Hindu Business Line, August 31, p 1
Anonymous (2011) Annual Report of AICRP (sunower)
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Basappa H (2007) Changing scenario of insect pests of
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Basappa H (2008) Biodiversity of arthropods and verte-
brates in sunower ecosystem. Paper presented in Dr.
Fig. 23.4 Cocoons of A. bambawalei Leslie C. Coleman memorial National Symposium on
Plant protection, UAS, Bangalore, 46 December 2008
Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
The fungal pathogen Verticillium lecanii was
Pseudococcidae and Putoidae) with data on geograph-
able to bring about 50 % reductions in the mealy- ical distribution, host plants, biology and economic
bug population. Biopesticides may be effective importance. Intercept Limited, Andover, 686 p
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of the mealybug, Phenacoccus solenopsis Tinsley
natural enemies (Basappa 2008).
(Homoptera: Pseudococcidae) infesting sunower
(Helianthus annuus L.). MSc. (Agri.) thesis in
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23.5.2 Under Field Conditions Agricultural Sciences, Bengaluru, 126 p
Chandrashekar, Jagadish KS, Shadakshari YG, Basan
Gowda G (2012) Ecology and host range of the mealy-
Profenophos 50 EC (0.05 %), methomyl 40 SP bug Phenacoccus solenopsis Tinsley (Hemiptera:
(0.04 %), dimethoate 30 EC (0.06 %) and dichlo- Pseudococcidae) in sunower eco-system. In:
rovos 76 WSC (0.15 %) were found to be most Proceedings of national seminar on Recent trends in
Pest management and conservation of our Natural
effective in controlling the mealybug on sunower
Enemies organized by the PG and Research
under eld conditions (Chandrashekar 2011; Department of Zoology, V.O. Chidambaram College,
Anonymous 2011). The insecticides can be used Thoothukudi-628008, from 1516 March 2012, p 3
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Gopalkrishanan N (2009) Natural occurrence of ento-
spray can be applied after 10 days of rst applica-
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tion if the pest population persists (Anonymous nopsis Tinsley in India. In: Proceedings of national
2011). Methomyl 40 SP (0.04 %) and profeno- symposium Bt Cotton: Opportunities and Prospects,
phos 50 EC (0.05 %) recorded maximum net CICR Nagpur, 121 p
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bambawalei Hayat (Hymenoptera: Encyrtidae), DOR Cotton mealybug, Phenacoccus solenopsis Tinsley- a
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Jagadish KS, Shadakshari YG, Jagannatha DP, Karuna K, Rathod PK, Mane PN, Lande GK, Sable YR (2008) First
Geetha KN, Puttarangaswamy KT (2008) Role of ant record of mealybug, Maconellicoccus hirsutus (Green)
(Hymenoptera: Formicidae) fauna in sunower on sunower in western Vidharbha (MS). Insect
(Helianthus annuus L.). Paper presented in Dr. Leslie Environ 14(1):18
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Pulses
24
S.K. Singh, S.D. Mohapatra, and P. Duraimurugan
Fig. 24.1 Mealybug damage to pigeon pea: (a) P. solenopsis on pigeon pea, (b) P. marginatus on pigeon pea and
(c) P. maraginatus on Phaseolus vulgaris
24.7 Management
References
24.7.1 Chemical Control
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tamethrin] (0.00125 %) caused 89.2, 88.1, 68.0,
Willink (Hemiptera: Pseudococcidae) in two ecologi-
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Soybean
25
M. Mani
Mealybugs are reported to be injurious to soy- feed on plant juices. These whitish crawlers are
bean in India, Kentucky (USA), Sri Lanka, seen attached to the roots (http://extension.entm.
Indonesia, and so forth (Table 25.1). purdue.edu/pestcrop/2011/issue14/index.html).
Pseudococcus sorghiellus has been found in Planococcus citri was known to suck the sap
association with elds that exhibit symptoms from stem at pod formation stage in Maharashtra,
similar to potassium deciency (yellowed leaf India (Jadhav et al. 1996). The development of
margins and stunted plants), often in elds that Pl. citri and the parasitoid Anagyrus pseudococci
recently hosted alfalfa. It seems that this mealy- (Girault) was better on soybeans than on
bug species is fairly common on other plant spe- Streptocarpus hybridus or Aeschynanthus ellipti-
cies, reported from Indiana, Illinois, and cus (Copland et al. 1993). The average develop-
Pennsylvania. The mealybug species is often mental periods of males and females of N.
found to be tended by ants, which eat the honey- vasatator on soybean were 15.46 and 14.62 days,
dew excreted by the mealybugs and in turn pro- respectively. The average preoviposition, ovipo-
tect the mealybugs from predators. Similar to sition, and fecundity were 7.66 days and 78.67
other mealybugs in appearance, these small, eggs on soybean (Saha and Ghosh 2001).
whitish insects live beneath the soil surface and
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Gimpel WF Jr, Miller DR (1996) Systematic analysis of Saha A, Ghosh AB (2001) Biological studies on the
the mealybugs in the Pseudococcus maritimus com- mealybug, Nipaecoccus viridis (Newstead) on various
plex (Homoptera: Pseudococcidae). Contrib Entomol host plants. Uttar Pradesh J Zool 21(1):7578
Int 2(1):163 Thippaiah M, Kumar NG (1999) Dysmicoccus sp.
Jadhav RG, Madane NP, Kathamale DK (1996) Record of (Pseudococcidae: Homoptera): a pest of soyabean in
soybean as a new host in India for citrus mealybug. Karnataka. Insect Environ 5(2):70
Insect Environ 2:90 Tooker J (2011) A new pest to consider: trochanter mealy-
Rajadurai S, Thyagarajan V (2003) Mulberry sap sucking bug. http://www.farms.com/FarmsPages/
pests. Indian Silk 42(4):58 Commentary/DetailedCommentary/tabid/192/
Srivastava OS (1972) Soybean, a new host record of Default.aspx?NewsID=43316
mealybug, Nipaecoccus vastator (Mask.) (Hemiptera: Williams DJ (2004) Mealybugs of southern Asia. The
Pseudococcidae) in India. Indian J Entomol Natural History Museum/Southdene SDN. BHD.,
34(3):251252 London/Kuala Lumpur, 896 p
Cotton
26
V.S. Nagrare, S. Kranthi, Rishi Kumar,
B. Dharajothi, M. Amutha, and K.R. Kranthi
Mealybugs are injurious to cotton in India, Mealybugs are small sap-sucking insects that cause
Pakistan, China, Brazil, Australia, Ethiopia, severe economic damage to cotton. They attack
USA, Emerald and so on. (Table 26.1). Among different parts of the growing cotton plant, namely
the mealybug species, the solenopsis mealybug main stem, branches and underdeveloped owers
Phenacoccus solenopsis Tinsley was found to that produce bolls of smaller size. The boll opening
cause severe economic damage in all nine major when adversely affected result in serious reduc-
cotton-growing states of India, resulting in loss tion in yield. Excretion of honeydew attracts ants
worth of US $1620 million while reducing and also contributes to the development of black
yields up to 50 % in the affected elds in 2007 sooty mould. Plants severely affected with sooty
(Nagrare et al. 2009). Paracoccus marginatus mould have the appearance of burn symptoms.
Williams and Granara de Willink also caused An infested cotton plant shows the symptoms
havoc on cotton around Coimbatore district of such as white uffy mass on underside of leaves,
Tamil Nadu. Besides these two mealybugs, near growing tips, along leaf veins and on stem,
Nipaecoccus viridis (Newstead), Maconellicoccus distorted or bushy shoots. Plants infested by
hirsutus (Green), Ferrisia virgata Cockerell, mealybugs during vegetative phase exhibit symp-
Ferrisia malvastra (Mc Daniel) in central India toms of distorted, bushy shoots, crinkled and/or
and Rastrococcus iceryoides (Green) in south twisted bunchy leaves and stunted plants that dry
India were also reported on cotton in traces. completely in severe cases.
Table 26.1 Mealybug species recorded on cotton in different regions of the world
Species Region/Country References
Ferrisia consobrina Australia, Ethiopia, Williams (2004)
Williams and Watson Neotropical and
Pacic region
Ferrisia malvastra (Mc Daniel) India Jhala et al. (2008), Suresh and Kavitha (2008a)
Ferrisia virgata Cockerell India Saminathan and Jayaraj (2001), Suresh (2008),
Vennila et al. (2010a), Anonymous (2010),
Nagrare et al. (2011), Tanwar et al. (2011),
Nagrare et al. (2012), Anonymous (2013b)
Brazil Torres et al. (2013)
Cambodia Williams (2004)
Maconellicoccus hirsutus (Green) India Misra (1920), Dhawan et al. (1980),
Muralidharan and Badaya (2000),
Suresh (2008), Nagrare et al. (2009),
Pinjarkar et al. (2009), Vennila et al. (2010b)
Nipaecoccus viridis (Newstead) India Misra (1920), Vennila et al. (2010b), Nagrare
et al. (2011), Thomas and Ramamurthy (2011)
Paracoccus burnerae (Brain) Ethiopian region Ben-Dov (1994)
Paracoccus marginatus India Anonymous (2008), Amutha et al. (2009),
Williams and Granara de Willink Banu et al. (2010), Dharajothi et al. (2010),
Tanwar et al. (2010),
Jeyarani et al. (2011), Nagrare et al. (2011),
Sakthivel et al. (2012)
Argentina Granara de Willink (2003)
Paracoccus solani Australia, Peru and Ben-Dov (1994)
Ezzat and McConnel Costa Rica
Phenacoccus madeirensis Green India Shylesha and Joshi (2012)
Phenacoccus solenopsis Tinsley China Wang et al. (2009), Wu and Zhang (2009),
Zhang et al. (2010)
India Dutt (2007), Hodgson et al. (2008), Jhala et al.
(2008), Thomas et al. (2008), Anonymous (2009),
Bhosle et al. (2009), Dhawan and Saini (2009),
Dhawan et al. (2009), Hanchinal et al. (2009),
Kumar et al. (2009a), Monga et al. (2009),
Nagrare et al. (2009), Saini et al. (2009),
Bisane et al. (2010), Suresh et al. (2010),
Vennila et al. (2010b), Hanchinal et al. (2011),
Kumar et al. (2011), Tanwar et al. (2011)
Pakistan Zaka et al. (2006), Muhammad (2007),
Hodgson et al. (2008), Abbas et al. (2009),
Afzal et al. (2009), Abbas (2010), Abbas et al.
(2005), Abbas et al. (2010), Ashfaq et al. (2010),
Khuhro et al. (2011), Sahito et al. (2011)
Brazil Silva (2012)
Ethiopia Anonymous (2013a)
USA Fuches et al. (1991)
Australia Admin (2010), Charleston and Murray (2010),
David and Charleston (2010), IPPC (2010)
Emerald and Burdekin Admin (2010)
Nigeria Akintola and Ande (2008)
Sri Lanka Prishanthini and Laxmi (2009)
(continued)
26 Cotton 273
correlated with temperature, whereas negative during the seventh meteorological week, which
correlation was observed with relative humidity coincides with the higher population of mealy-
and rainfall (Dhawan et al. 2009). bug. Mealybug population was signicantly and
It appeared that high rainfall has washed away positively correlated with maximum temperature
all the small crawlers. Moreover, the high rainfall (0.775) and negatively correlated with other
has favoured the growth of entomopathogens on parameters. Among predators, chrysopids were
the mealybugs. It is evident that enough humidity signicantly correlated with relative humidity
favours the multiplication, but the intense rainfall (0.289) and others were non-signicant. The
adversely affects the spread and reduces the mealybug parasitoid cocoons were positively
intensity (Jeyakumar et al. 2009). In Raichur, correlated with maximum temperature (0.421)
Karnataka, India, mealybug infestation started but negatively correlated with other meteorologi-
appearing in September and gradually increased cal parameters (Hanchinal et al. 2010).
as the crop growth advanced. The population was
0.50/10-cm apical shoot in the 38th meteorologi-
cal week and progressively increased throughout 26.5 Natural Enemies
the season. The population reached to 115.42/10-
cm apical shoot in the third week of January and 26.5.1 P. solenopsis
thereafter increased suddenly to reach 180.42/10-
cm apical shoot in the seventh meteorological 26.5.1.1 Parasitoids
week. Later on, the infestation of mealybug The encyrtid parasitoid Aenasius bambawalei
declined gradually and reached to 146.64/10-cm Hayat was recorded on Ph. solenopsis-infesting
apical shoot in the 14th meteorological week. In cotton and other crops in India (Hayat 2009). It
general, predator population was low during the was reported from north and central India
cropping season. Maximum population of cocci- (Tanwar et al. 2011), Haryana (Ram Pala et al.
nellids, chrysopids and spiders were 0.14, 0.13 2009), Punjab (Dhawan et al. 2011), Tamil Nadu
and 0.16 per plant, respectively, during the sea- (Sankar et al. 2011; Amutha et al. 2009),
son. Parasitoid cocoons ranged between 0.52 and Karnataka (Hanchinal et al. 2009), all in India,
20.02 %. The activity of the parasitoid Aenasius Pakistan (Arif et al. 2011a) and China (Chen
bambawalei started during the 44th meteorologi- Hua-Yan et al. 2010). The parasitoid seems host
cal week and later on increased gradually to reach specic, having excellent searching ability in
the peak during seventh to ninth meteorological attacking mealybugs in colonies as well as those
weeks. Highest parasitoid (20.65 %) was recorded present solitarily on different host plants of
26 Cotton 275
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bug on cotton elds in the States of Bahia and Paraba, Nipaecoccus viridis (Newstead) (Hemiptera:
Brazil. Bragantia Campinas 71(4):467470 Pseudococcidae). Munis Entomol Zool 6(1):455459
Solangi GS, Mahmood R (2011) Biology, host specicity Thomas A, Chaudhary B, Singh J, Ramamurthy VV
and population trends of Aenasius bambawalei Hayat (2008) Preliminary observations on the non-target
and its role in controlling mealy bug Phencoccus sole- insect communities in Bt cotton. Indian J Entomol
nopsis Tinsley at Tandojam Sindh. Paper presented in 70(4):400402
5th ICAC international cotton advisory committee Torres CS, Oliveira MD, Torres JB (2013) Host selection
held at Lahore, Pakistan, 2325 February and establishment of striped mealybug, Ferrisia vir-
Solangi GS, Lohar MK, Abro GH, Buriro AS (2012) gata, on cotton cultivars. Phytoparasitica 41(1):3140
Biology and release of exotic predator Cryptolaemus Vennila S, Deshmukh AJ, Pinjarkar D, Agarwal M,
montrouzieri Mulsant on mealybug Phenacoccus sole- Ramamurthy VV, Joshi S, Kranthi KR, Bambawale
nopsis Tinsley at Tandojam. Sarhad J Agric OM (2010a) Biology of the mealybug, Phenacoccus
28(3):429435 solenopsis on cotton in the laboratory. J Insect Sci
Suganthi A, Kalyanasundaram M, Mahalaksmi V (2009) 10:115119
A buttery predator on cotton mealy bugs. Online edi- Vennila S, Ramamurthy VV, Deshmukh AJ, Pinjarkar DB,
tion of Indias National Newspaper The Hindu Agarwal M, Pagar PC, Prasad YG, Prabhakar M,
Thursday, November 12 Kranthi KR, Bambawale OM (2010b) A treatise on
Suresh S (2008) Cotton mealybug complex. In Proceedings mealybugs of Central Indian cotton production sys-
of the National Consultation on Mealybug tem, Technical bulletin no 24. National Centre for
Management. Central Institute for cotton Research, Integrated Pest Management, New Delhi, 39p
2829 January, Nagpur, India, pp 1517 Wang YP, Wu SA, Zhang RZ (2009) Pest risk analysis of
Suresh S, Kavitha PC (2008a) New records of Coccoidea a new invasive pest Phenacoccus solenopsis, to China.
in India. In: Branco M, Franco JC, Hodgson CJ (eds) (in Chinese; Summary in English). Chin Bull Entomol
Proceedings of the XI international symposium on 46(1):101106
scale insect studies, Oeiras, Portugal, 2427 September Williams DJ (2004) Mealybugs of Southern Asia. The
2007. ISA Press, Lisbon, p 322 natural history museum, London, and Southdene.
Suresh S, Kavitha PC (2008b) Seasonal incidence of eco- SDN BHD, Kuala Lumpur, 896 pp
nomically important coccid pests in Tamil Nadu. 285 Wu SA, Zhang RZ (2009) A new invasive pest,
291 In: Branco M, Franco JC, Hodgson CJ (eds) Phenacoccus solenopsis threatening seriously to cot-
Proceedings of the XI international symposium on ton production. Chin Bull Entomol 46(1):159162
scale insect studies, Oeiras, Portugal, 2427 September Zaka SM, Saeed S, Bukhari SA, Baksh E (2006)
2007. ISA Press, Lisbon, 322 p Mealybug, Phenacoccus solenopsis (Homoptera:
Suresh S, Jothimani R, Sivasubrmanian P, Karuppuchamy Pseudococcidae): a novel pest of cotton in Pakistan.
P, Samiyappan R, Jonathan EI (2010) Invasive mealy- In: Proceedings of 34th Pakistan (SAARC) countries
bugs of Tamil Nadu and their management. Karnataka science conference, University of Veterinary and
J Agric Sci 23(1):69 Animal Sciences, 810 February 2006. Lahore,
Surulivelu T, Sonai Rajan T, Dharajothi B (2010) Pakistan, p 32
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(Paracoccus marginatus) in cotton. CICR Newsl invasive mealybug, Phenacoccus solenopsis Tinsley
6(1):7 (Hemiptera: Pseudococcidae) in China. In: Potential
Tanwar RK, Jeyakumar P, Vennila S (2010) Papaya invasive pests workshop held at Miami, Florida, 1014
mealybug and its management strategies, Technical October 2010, 71p
Jute and Allied Fibre Crops
27
M. Mani and S. Satpathy
27.2 Damage
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India The mealybug infestation in India went up to
e-mail: mmani1949@yahoo.co.in 80 % in case of jute with average intensity of
S. Satpathy 23 in 04 scale. The plant infestation in mesta
Central Research Institute for Jute and Allied Fibres, was 60 % with average intensity of 4 in the
Barrackpore, Kolkata 700120, India 04 scale.
e-mail: satpathy.iivr@gmail.com
Table 27.1 List of mealybugs recorded on Kapok in dif- In M. hirsutus-infested roselle (Hibiscus sab-
ferent countries
dariffa var. altissima) plants, the number of pods
Mealybug Species Country Reference averaged 13.43/plant, the number of seeds 10.57/
Deltococcus tafaensis Ghana Ben-Dov pod and the percentage germination of the seed
(Strickland) (1994)
78.61, as compared with 24.40 pods/plant, 27.83
Maconellicoccus hirsutus Thailand Williams
seeds/pod and 87.63 % germination in uninfested
(Green) (2004)
Planococcus citri (Risso) Ben-Dov
plants (Ghose 1971). Maconellicoccus hirsutus is
(1994) emerging to be a major pest of mesta, particularly
Planococcus indicus India, Ben-Dov in the peninsular India. It causes bunchy top
(Avasthy & Shafee) China (1994) which is a serious malady of mesta crop. The
Planococcoides Ben-Dov bre crops Hibiscus sabdariffa var. altissima
nijalensis (Laing) (1994) (roselle), H. Cannabinus and Boehemeria nivea
Parcoccua marginatus India Mani et al have been reported to be the major hosts of
Williams and Granara de (2012)
Willink mealybugs in West Bengal, India, and Bangladesh
Planococcus lilacinus Indonesia Williams (Ghose 1972a; Singh and Ghosh 1970). The
(Cockrell) (2004) reduction in bre yield of roselle is to the extent
Rasrococcus iceryoides Thailand Williams of 21.4 % (Ghose 1971) to 40 % (Raju et al.
(Green) (2004) 1988). The salivary toxin injected during feeding
27 Jute and Allied Fibre Crops 285
causes characteristics distortion of leaves, short- against M. hirsutus because of its habit of feeding
ening if internodes and bushy top symptom in inaccessible parts and waxy covering of the
(Singh and Ghosh 1970; Ghose 1972b; Williams body (Williams 1996). Any pesticide used against
1996). Infestation by mealybug, M. hirsutus, M. hirsutus should be carefully selected to avoid
reduces the linear growth of the stem and petiole injury to its natural enemies, since they are likely
and markedly reduces the size, distort the mesta to be important in helping to keep populations at
leaves, causing bunchy top symptoms (Dutta low levels in the long term. The farmers used
et al. 1951), causing 1520 % reduction in bre wide array of insecticides without any apprecia-
yield (Das and Singh 1986). Its infestation causes ble result against Ph. solenopsis. On the other
damage to both bre and seed crop. Ferrisia vir- hand, insecticide spray suppressed the natural
gata has also been reported to induce stiffness of enemy activity, resulting in build-up of the
the ramie plants which makes the extraction of mealybug population (Satpathy et al. 2009).
bres by machine very difcult. In Bangladesh, it Cryptolaemus montrouzieri is a very good candi-
caused 6570 % infestation of mesta plants (Jalil date for the biological control of M. hirsutus and
and Kabir 1971). There was an outbreak of Ph. F. virgata, and this predator can be utilised for the
solenopsis in jute and mesta in West Bengal. The suppression of mealybugs. Acerophagus papayae
plant infestation in jute (cv. JRO524) was 6080 can be utilised for the control of P. marginatus
% with an average intensity of 23 in 04 scale, and so also Aenasius bombavale for Ph. solenop-
while it was 60 % and 40 %, respectively, in the sis in bre crops.
case of mesta (cv Local). Analysis of weather
factors indicated that the warm and dry weather
condition during summer might be the predispos- References
ing factor for the mealy outbreak in jute and
mesta (Satpathy et al. 2009). Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
Pseudococcidae and Putoidae) with data on geograph-
ical distribution, host plants, biology and economic
27.3 Natural Enemies importance. Intercept Limited, Andover, 686 p
Das LK, Singh B (1986) Economic control of
Maconellicoccus hirsutus (Green) infesting mesta.
The coccinellid predators Brumoides suturalis
Indian J Agric Sci 56(5):373375
(F.) and Scymnus nubilus (Muls.) and the encyr- David BV, Ananthakrishnan TN (2004) General and
tid parasite Anagyrus sp. are the important natu- applied entomology. Tata McGraw-Hill Publishing,
ral enemies on M. hirsutus. Six species of ants New Delhi, 1184p
Dutta N, Mukherjee PK, Gupta NS (1951) Preliminary
were found attending M. hirsutus in West Bengal,
observations on the incidence of Phenococcus hirsu-
India (Ghose 1970a). In Andhra Pradesh, India, tus Gr. and its effect on the growth of H. subdariffa
Spalgis epeus (Westwood), Hyperaspis maindroni Lin.var. altissima Hort. India. Indian J Agric Sci
Sicard, Autoba silicula Swinhoe and 21:231237
Galanihe LD, Watson GW (2012) Identication of
Brinckochrysa scelestes Banks were found prey-
Rastrococcus rubellus Williams (Hemiptera:
ing on eggs and nymphs of M. hirsutus, an impor- Pseudococcidae) on mango: a new record to Sri Lanka.
tant pest of kenaf (mesta) (Rao et al. 1984). Tropical Agric Res Ext 15(2):710
Ghose SK (1961) Studies on some coccids of economic
importance of West Bengal, India. Indian Agric
5:5778
27.4 Management Ghose SK (1970a) Predators, parasites and attending ants
of the mealybug, Maconellicoccus hirsutus (Green)
Sprays of methyl demeton at 0.2 % were found (Pseudococcidae, Hemiptera). Plant Protect Bull India
22(3):2230
highly effective in controlling populations of M.
Ghose SK (1970b) Chemical control of the mealybug,
hirsutus on roselle Hibiscus sabdariffa (Ghose Maconellicoccus hirsutus (Green) (Pseudococcidae,
1970b). The chemicals have limited effectiveness Hemiptera). Plant Protect Bull India 22(3):1721
286 M. Mani and S. Satpathy
Ghose SK (1971) Assessment of loss in yield of seeds of Rao PRM, Kanakaraju A, Apparao RV, Azam KM (1984)
roselle (Hibiscus sabdariffa L. var. altissima Wester) Predators on mealy bug of mesta. Q Newsl FAO Asia
due to the mealybug, Maconellicoccus hirsutus Pac Plant Protect Commission 27(4):12
(Green) (Pseudococcidae: Hemiptera). Indian J Agric Ritchie AH (1935) Report of the entomologist. Tanganyika
Sci 41(4):360362 Department of Agriculture Report for 1934, pp 7383
Ghose SK (1972a) Biology of the mealybug Satpathy S, Gotyal BS, Ramasubramanian T,
Maconellicoccus hirsutus (Green) (Pseudococcidae, Bhattacharyya SK, Laha SK (2009) Mealybug infesta-
Hemiptera). Indian Agric 16(4):232332 tion in jute and mesta crop a case study. In: National
Ghose SK (1972b) Morpho-histological changes in some symposium on climate change, plant protection, and
economic plants due to the infestation of mealy bug, food security interface, West Bengal, Kalyani, 1719
Maconellicoccus hirsutus (green) (Hemiptera: December 2009, p 78
Pseudococcidae). Indian J Agric Sci 42(4):329334 Singh MP, Ghosh SN (1970) Studies on Maconellicoccus
Hall WJ (1921) The Hibiscus mealybug (Phenacoccus (Phenococcus) hirsutus Gr. causing bunchy top in
hirsutus Green). Bull Minis Agric Egypt 17:128 mesta. Indian J Sci Indus 4(2):99105
Jalil AFMA, Kabir AKMF (1971) Some studies on Tanwar RK, Jeyakumar P, Vennila S (2010) Papaya
Pseudococcus virgatus Ckll. infesting Mesta and mealybug and its management strategies. Technical
Kenaf in East Pakistan. Agric Pak 22(2):237240 Bulletin 22, National Centre for Integrated Pest
Kundu BC, Basak KC, Sirkar PB (1959) Jute in India- a Management, New Delhi, 22p
monograph. Indian Central Jute Committee, Calcutta, Tripathy RL, Ram S (1971) A review of entomological
pp 159170 researches on Jute, Mesta. Sunnhemp and allied
Mani M, Shivaraju C, Sylesha AN (2012) Paracoccus bres. ICAR technical bulletin no. 36. New Delhi:
marginatus, an invasive mealybug of papaya and its Indian Council of Agricultural Research
biological control. J Biol Control 26(3):201216 Williams DJ (1996) A brief account of pink hibiscus
Pollard GV (1995) Pink hibiscus mealy bug in the mealybug, Maconellicoccus hirsutus (Hemiptera,
Caribbean. CARAPHIN News 12:12 Pseudococcidae), a pest of agriculture and horticulture
Raju AK, Rao PRM, Apparao RV, Reddy AS, Rao KKP with description of two related species from Southern
(1988) Note on estimation of losses in yield of mesta Asia. Bull Ento Res 86:617628
due to mealy bug, Maconellicoccus hirsutus Green. Williams DJ (2004) Mealybugs of southern Asia. The
Jute Dev J 8(1):3435 Natural History Museum/Southdene SDN. BHD,
London/Kuala Lumpur, 896 p
Sugarcane
28
R. Jayanthi, J. Srikanth, and S.N. Sushil
28.1 Mealybug Species chari was reported for the rst time in Cuba along
with observations on three other species, includ-
Mealybugs are widespread throughout the ing S. sacchari (Williams et al. 2001).
sugarcane-growing tracts of the world but all of Kiritshenkella sacchari (Fig. 28.1bc) and
them seldom attained major pest status (Tohamy Antonina graminis are also commonly encoun-
et al. 2008). According to Williams (1970), 30 tered in Tamil Nadu, India. The incidence pattern
species of mealybugs are known to attack sugar- of these three species revealed that K. sacchari
cane in different regions of the world (Table has the potential to emerge as an important pest,
28.1). Nine species of mealybugs are known to especially under drought conditions. While
occur in India. A detailed compendium on the Dysmicoccus carens (Fig. 28.1de) was observed
Indian species of sugarcane mealybugs with infesting the foliage of sugarcane hybrids,
respect to their biology, factors inuencing their Pseudococcus sp. was observed on sugarcane
buildup, crop damage, and management aspects rootlets (Jayanthi et al. 1995). Pseudococcus sac-
has been documented by Jayanthi (1986). The charicola was reported from south Andaman on
pink mealybug Saccharicoccus sacchari sugarcane leaves (Veenakumari and Mohanraj
(Fig. 28.1a) is the most ubiquitous species in 1995). Dysmicoccus carens was recorded in
India. Outside India, S. sacchari on sugarcane Andhra Pradesh by Rao et al. (2008).
was reported from Alexandria and Egypt (Hafez
and Salama 1969; Mesbah et al. 1976). It may be
native to Eastern Africa but was also reported to 28.2 Damage
occur in Formosa, Malaysia, Philippines, Java,
Hawaii, Samoa, Australia, Syria, Egypt, Madeira, Nymphs and adults suck the sap from leaves,
Argentina, Peru, British Guiana, Mexico, nodes, and internodes of canes. Severe infesta-
Caribbean Islands, Mauritius, South Africa and tion results in yellowing of leaves, stunting of
East Africa (Clausen 1978). Kiritshenkella sac- canes and poor germination in the case of S. sac-
chari attack.
R. Jayanthi (*) J. Srikanth Loss of sap may kill the young shoots in the
ICAR-Sugarcane Breeding Institute, case of Ps. saccharicola or may result in a marked
Coimbatore 641007, India setback in cane growth, ultimately leading to
e-mail: jayajanani21@yahoo.com total drying of the crop as evident in instances of
S.N. Sushil Ph. saccharifolii infestation. The stalks on which
ICAR-Indian Institute of Sugarcane Research, the mealybug colonies have fed and perished can
Lucknow 226002, India
Table 28.1 List of mealybugs recorded on sugarcane in different regions of the world
Mealybug species Region/Country Reference
Antonina graminis (Maskell) Williams (1970)
Hawaii Pemberton (1938)
India Ahmad (1942)
Sri Lanka Kumarasinghe (2003)
Texas Riherd (1950)
Cannococcus ikshu Williams and Papua New Guinea Ben-Dov (1994)
Watson
Chorizococcus rostellum (Lobdell) Williams (1970)
Chorizococcus talipikanus Williams Papua New Guinea Ben-Dov (1994)
and Watson
Dysmicoccus boninsis (Kuwana) Indonesia and Sri Lanka Williams (2004)
Dysmicoccus brevipes (Cockerell) India Williams (2004)
Dysmicoccus carens sp.n. India Williams (1970)
Dysmicoccus cryptus (Hempel) Williams (1970)
Dysmicoccus trispinosus (Hall) Williams (1970)
Eumyrmococcus smithii Silvestri China and Japan Williams (2004)
Exallomochlus hispidus (Morrison) Indonesia, Java and Malaysia Williams (2004)
Formicococcus lingnani (Ferris) Indonesia Williams (2004)
Ferrisia virgata (Cockerell) Williams (1970)
Heliococcus summervillei Brookes Australia and Pakistan Ben-Dov (1994)
Kiritshenkella sacchari (Green) [= Cuba Williams et al. (2001)
Ripersia sacchari (Green)] India Ayyar (1919)
Pakistan Ali (1995)
Bangladesh, Burma, India, and Williams (2004)
Pakistan
Maconellicoccus hirsutus (Green) Williams (1970)
Madagasia cincinnata sp.n. Williams (1970)
Mizococcus sacchari Takahashi Williams (1970)
Taiwan Williams (2004)
Neoripersia ogasawarensis (Kuwana) Ogasawara Islands Ben-Dov (1994)
Paracoccus eastopi Williams Nigeria Williams (1970); Williams (2004)
Paracoccus spinulosus (De Lotto) Williams (1970)
Uganda Ben-Dov (1994)
Phenacoccus hargreavesi (Laing) Williams (1970)
Ethiopia Ben-Dov (1994)
Phenacoccus parvus Morrison Ethiopian, Neotropical, and Pacic Ben-Dov (1994)
region
Phenacoccus saccharifolii (Green) Williams (1970)
India Green (1908), Isaac and Misra
(1933)
India, Nepal, and Pakistan Williams (2004)
Planococcus citri (Risso) Williams (1970)
Planococcoides lindingeri Egypt and Israel Ben-Dov (1994)
(Bodenheimer)
Planococcoides lingnani (Ferris) China Ben-Dov (1994)
Planococcus minor (Maskell) Trinidad Francis et al. (2012)
(continued)
28 Sugarcane 289
be recognized by residues of wax, honeydew, and juice signicantly in cv. Co 1148 (Atiqui and
sooty mold that persist for months (Jayanthi Murad 1992).
1986.
Severe infestation of S. saccchari in canes
reduced sucrose content by 24.1 % and brix by 28.3 Factors Inuencing Buildup
16.2 % (Kalra and Sidhu 1964). Varietal reaction
in terms of infestation levels and yield and qual- Agroclimatic conditions and crop management
ity parameters due to S. sacchari infestation have factors are known to have impact on the mealy-
been elucidated by Jayanthi (1991). In Uttar bug abundance, both positively and negatively.
Pradesh, India, S. sacchari infestation reduced While dry conditions generally favor mealybugs,
sugar brix, sucrose, purity, and available sugar rainfall exerts a negative inuence, apparently by
content by 10.64, 16.44, 6.14, and 12.92 %, dislodging colonies in detrashed crop and pro-
respectively but did not affect the volume of cane moting the growth of entomopathogenic fungi.
290 R. Jayanthi et al.
However, Ph. saccharifolii was observed to relation with incidence in Andhra Pradesh, India.
multiply rapidly with the onset of monsoon Water stress conditions, small dry spells,
(Mohammad Ali 1962). In general, conditions neglected ratoons and repeated ratooning
that are least favorable for the vegetative growth enhanced infestation (Rao et al. 2009). Mealybug
of the sugarcane plant appear to be most favor- abundance is also known to be inuenced by
able for the increase of S. sacchari. Reduction in plant factors such as the nature of leaf sheath.
S. sacchari populations was observed with Varieties with loose clasping leaf sheath harbor
enhanced levels of irrigation in drip-irrigated higher levels of mealybug populations than those
plots (Parsana et al. 1994). Ratoons were not sus- with tight clasping leaf sheath. Self-stripping
ceptible as a general rule; on the other hand, the varieties are less prone to mealybug infestation.
plant crop of some commercial hybrids was more According to Jayanthi (1991) and Jayanthi et al.
susceptible than the ratoon counterparts (Jayanthi (1994), stem hardness was not found to inuence
1991). Saccharicoccus sacchari outbreak colonization by S. sacchari; some biochemical
occurred in years of moderate temperature during parameters were related to infestation by S. sac-
MarchApril, and temperature had a positive cor- chari. Ants, generally found associated with
28 Sugarcane 291
mealybugs, have been shown to interfere with Mealybug-tolerant clones were identied in a
predator activity (Srikanth et al. 2001), thereby series of other screening studies with clones and
possibly enhancing the mealybug density. standard varieties at different locations (Abdullah
2009; Abdullah et al. 2006a, b, 2010). In studies
with 43 germplasms against S. sacchari in Egypt,
28.4 Varietal Susceptibility C 46-117, Co 237, Co 290, Co 997, CP 31-294,
CP 34-38 and CP 52-43 were classied as resis-
No truly resistant varieties are available because tant (Solouma 2002). The varieties Giza 96/74
mealybugs are capable of infesting any sugarcane and Ph 8013 were less susceptible to S. sacchari
genotype. However, higher levels of infestation based on percent infested internodes and number
are often observed in genotypes with loose clasp- of mealybugs per stalk (Tohamy et al. 2008).
ing leaf sheath. The commercial varieties CoC
671 and CoA 7601 with tight clasping leaf sheath
always registered low incidence of S. sacchari 28.5 Natural Enemies
(Sithanantham 1973; Mehta et al. 1981). The
genotypes Co 740, Co 6806, Co 8014, CoC 671, About 16 parasitoids, 13 predators and the
and CoA 7601 showed tolerance to S. sacchari entomopathogenic fungus Aspergillus parasiti-
(Anonymous 1992). In Assam, the varieties cus were recorded in different parts of India
CoBLN 9101, 9102, 9103, Co 6806, C0Jor-1, (Jayanthi 1986). The encyrtid Anagyrus punct-
C0Jor-2 and Co 740 were relatively resistant to S. ulatus Agarwal was found to be the most impor-
sacchari (Borah and Dutta 1995). Although none tant parasitoid in regulating the pest population
of the 24 sugarcane varieties screened were free in Gujarat; the parasitoid showed positive
from nymphs of S. sacchari, lowest densities of results in augmentative studies (Kapadia et al.
nymphs/internode were found on CoN 84136 and 1995). The activity of the parasitoid was noticed
CcN 84134, while the highest were on Co 87004 to be highest in July and lowest during
(Parsana et al. 1995). Similarly, none of the 17 NovemberDecember (Parsana et al. 1996). In
genotypes evaluated against borers and sucking Maharashtra, Chilocorus nigrita (F.) was found
pests in Maharashtra, India, were found to be attacking S. sacchari (Dorge et al. 1972). In
resistant to S. sacchari (Hole et al. 2009). The Uttar Pradesh, six parasitoids and four preda-
sugarcane cultivars Q 63 and Co 6501, catego- tors, including Batrachedra sp. near psilopa
rized as being lightly infested by S. sacchari and Meyrick (Lepidoptera: Momphidae), were
scale, showed higher quantities of phenols at har- reported on S. sacchari (Nigam 1983; Singh
vest compared to the other heavily infested culti- et al. 1997). The cecidomyiid predator
vars, namely Co 671, Co 6806, Co 740, and G Dicrodiplosis sp. was also recorded on S. sac-
229 (Jayanthi and Goud 2001). Differential bio- chari in Andhra Pradesh (Reddy and Aziz
logical parameters of D. carens observed on the 2000). The natural enemies of S. sacchari found
sugarcane genotypes Co 740, Co 7704, C 6806, in the neighboring Sri Lanka were listed by
CoC 671, and Co 6907 under laboratory condi- Rajendra (1974). Of these, the predatory droso-
tions indicated differential susceptibility of the philid Gitonides (Gitona) perspicax Knab, lar-
genotypes to the mealybug (Razak et al. 1994). vae of the nitidulid Carpophilus marginellus
Nine promising clones and a commercial cultivar Motsch. and ve encyrtid parasitoids of a genus
were considered susceptible to S. sacchari in near to Microterys were important in control-
Bangladesh. Lower infestation levels in clones I ling mealybug populations in the eld. The rat
155-91 and I 209-91 indicated that these might be Millardia meltada meltada gnawed through the
chosen as promising material to develop com- lower dry leaf sheaths and devoured the mealy-
mercial cultivars (Taleb and Rahman 2004). bugs at the nodes (Rajendra 1974).
292 R. Jayanthi et al.
Somalia (East Africa) as it failed to penetrate (Hemiptera: Pseudococcidae) attacking sugar cane in
Egypt. Sugar Cane Int 5:2426
under leaf sheaths where the mealybugs congre-
Abdullah M (2009) Incidence of some major insect pests
gate (Chairamonte 1933). In Hawaii, some con- to sugarcane clones in Bangladesh. Indian Sugar
trol of S. sacchari was achieved with its 59(3):1726
introduction in 1893 (Pemberton 1948, 1964). Abdullah M, Rahman MA, Biswas MM (2006a) Reaction
of some promising sugarcane clones to major insect
When C. montrouzieri was introduced with two
pests under eld condition. Indian Sugar
other predatory coccinellids, namely Hyperaspis 55(10):3742
sp. and Nephus sp., from India in 19681969, Abdullah M, Biswas MM, Siddiquee MNA (2006b)
and the encyrtid Anagyrus saccharicola Response of some promising sugarcane clones to
major insect pests of sugarcane. Indian Sugar
Timberlake (Hymenoptera: Encyrtidae) was
56(3):2328
introduced from East Africa in 1970 against S. Abdullah M, Alam MA, Rahman MA (2010) Tolerance
sacchari in Barbados, only the parasitoid had status of commercial varieties to major insect pests of
been recovered. Rapid spread, aided by addi- sugarcane. Indian Sugar 59(10):3138
Ahmad T (1942) Report of the second entomologist
tional releases, led to 8.39.7 % parasitism in
(Dipterist) in charge of the scheme for research on
differential rainfall areas by 1972 (Alam 1972). insect pests of sugarcane. Scientic Report of the
Interference by the attendant ant Camponotus Imperial Agricultural Research Institute, New Delhi,
compressus F. (Fig. 28.6f) through physical for 19401941, pp 6465
Alam MM (1972) The establishment of Anagyrus sac-
removal of stages of C. montrouzieri (Srikanth
charicola Timb. (Hymenoptera: Encyrtidae) in
et al. 2001) could be one of the reasons for the Barbados, West Indies, against the sugarcane mealy-
predators limited success. Anagyrus sacchari- bug, Saccharicoccus sacchari (Ckll.)
cola Timberlake releases against S. sacchari in (Hemiptera:Coccidae). Entomophaga 17(4):357363
Ali M (1995) Bionomics of sugarcane mealybug,
ve governorates of upper Egypt during 1999
Kiritshenkella sacchari (Green) (Homoptera,
2000 led to the parasitoids rapid establishment Pseudococcidae). Pakistan J Zool 27(1):1519
and spread with considerable increase in the Anonymous (1912) The sugarcane mealybug in Costa
rates of parasitism (Abd-Rabou 2002; Tohamy Rica. Bol Fom San Jose, Costa Rica. 7:466469
Anonymous (1992) Sugarcane coccoids. Sugarcane Breed
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(Homoptera: Pseudococcidae) on sugarcane, Williams DJ (1970) The mealybugs (Homoptera,
Saccharum ofcinarum Linnaeus a new record from Coccoidea, Pseudococcidae) of sugar-cane, rice and
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Pseudococcidae) in the British Virgin Islands: rst London/Kuala Lumpur, 896 p
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(Coleoptera: Coccinellidae). Proc Entomol Soc Wash cussion of a new record for the New World (Hem.,
112(4):565575 Pseudococcidae). Entomol Mon Mag
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Fruit Crops: Apple
29
M. Mani
Table 29.1 List of mealybug species infecting apple in different regions of the world
Mealybug species Region/Country Reference
Pseudococcus maritimus (Ehrhorn) New Zealand McKenzie (1972)
Pseudococcus fragilis Brain, South Africa Myburgh et al. (1975), Swart
Pseudococcus obscurus Essig. (1977)
Pseudococcus viburni (Signoret) South Africa Stokwe and Malan (2010)
France Kreiter et al. (2005)
New Zealand Charles et al. (2004)
Pseudococcus longispinus (Targioni South Africa Stokwe and Malan (2010)
Tozzetti) Japan Morimoto (1976)
Pseudococcus calceolariae (Maskell) South Africa Stokwe and Malan (2010)
Japan Morimoto (1976)
Pseudococcus comstocki (Kuwana) Japan Morimoto (1976)
New York and California
Planococcus citri (Risso) Florida
Phenacoccus aceris Sig. Nova Scotia Gilliatt (1935/1936)
British Columbia Madsen and Proctor (1982),
Marshall and Pickett (1944);
Marshall (1953), Kozar et al.
(1989)
Dysmicoccus debregeasiae (Green) Bangladesh Williams (2004)
Maconellicoccus hirsutus (Green) Ben-Dov (1994)
Phenacoccus mespili (Signoret) France and Russia Ben-Dov (1994)
Phenacoccus graminicola Leonardi Australia and New Zealand Ben-Dov (1994)
including exclusion cages, limb-banding, and Gilliatt FC (1936) Observations on the mealybug,
Phenacoccus aceris Sig. Can Entomol 68:133
visual inspection of shoots and fruits. The com-
Gilliatt FC (1939) The life history of Allotropa utlilis
plex of native natural enemies (which included Mues., a hymentoperous parasite of the orchard mealy
two encyrtid parasitoids, namely Pseudaphycus bug in Nova Scotia. Can Entomol 71:160161
websteri Timberlake and Mayridia sp.), a cocci- Grasswitz TR, Burts EC (1995) Effect of native natural
enemies on the population dynamics of grape mealy-
nellid beetle (Hyperaspis lateralis Mulsant), and
bug Pseudococcus maritimus (Homop; Pseducoccidae)
a chamaemyiid y (Leucopis verticalis Malloch) in apple and pear orchards. Entomophaga
provided a reasonably good control in orchards 45(1):105117
that had not been treated with insecticides for 12 Kozar F, Humble LM, Foottit RG, Orvos IS (1989) New
and little known scale insect (Homoptera: Coccoidea)
years. However, surveys indicated that most of
from British Columbia. J Entomol Soc BC 86:7077
these species were absent from orchards regu- Kreiter P, Delvare G, Giug L, Thaon M, Viaut M (2005)
larly sprayed with pesticides (Grasswitz and First inventory of the natural enemies of Pseudococcus
Burts 1995). viburni (Hemiptera, Pseudococcidae). [French]. Bull
Soc Entomol Fr 110(2):161164
Madsen HF, Proctor PJ (1982) Insects and mites of tree
fruits in British Columbia. Ministry of Agriculture and
References Food, Victoria, pp 170
Marshall J (1953) A decade of pest control in British
Bell VA, Walker JTS, Suckling DM, Manning LA, El Columbia. Proc Entomol Soc BC 49:711
Sayed AM, Shaw PW, Wallis DR, Miller JG, Shaw PV Marshall J, Pickett AD (1944) The present status of the
(2006) Trapping obscure mealybug (Pseudococcus apple mealybug, Phenacoccus aceris Sig., in
viburni) and its natural enemy Pseudaphycus maculip- British Columbia and Nova Scotia [Note]. Can
ennis (Hymenoptera: Encyrtidae) in apple orchards Entomol 76:19
(Poster Abstract). N Z Plant Protect 59:64 McKenzie DJ (1972) Control of codling moth, leaf roller
Ben-Dov Y (1994) A systematic catalogue of the mealy- and mealybug in apples. Orchardist N Z 46(5):155
bugs of the world (Insecta: Homoptera: Coccoidea: Morimoto R (1976) A study of the biological control of
Pseudococcidae and Putoidae) with data on geograph- Comstock mealybugs, Pseudococcus comstocki
ical distribution, host plants, biology and economic Kuwana. Rev Plant Prot Res 9:7289
importance. Intercept Limited, Andover. 686 p Muesbeck CFW (1939) A new mealybug parasite
Charles JG (1989) Pseudococcidae, mealybugs (Hymenoptera: Scelionidae). Can Entomol
(Homoptera). In: Cameron PJ et al (eds) A review of 71:158160
biological control of invertebrate pests and weeds in Myburgh AC, Rust DJ, Stubbings D (1975) Mealybugs on
New Zealand 18741987, Technical Communication apples and pears. Deciduous Fruit Grower
10. CAB International, Wallingford, pp 223236 25(7):176179
Charles JG (2001) Introduction of a parasitoid for mealy- Raine J, McMullen RD, Forbes AR (1986) Transmission
bug control: a case study under new environmental of the agent causing little cherry disease by the apple
legislation. N Z Plant Prot 54:3741 mealybug Phenacoccus aceris and the dodder Cuscuta
Charles JG, Allan DJ, Rogers DJ, Cole LM, Shaw PW, lupuliformis. Can J Plant Pathol 8:611
Wallis DR (2004) Mass-rearing, establishment and Stokwe NF, Malan AP (2010) Potential use of entomo-
dispersal of Pseudaphycus maculipennis, a biocontrol pathogenic nematodes for biological control of mealy-
agent for obscure mealybug. N Z Plant Prot bugs on apples and pears. SA Fruit J 9(3):3839, 42
57:177182 Swart PL (1977) Mealybugs: more economic manage-
Eastwell KC, Bernardy MG (2001) Partial characteriza- ment on apples and pears. Deciduous Fruit Grower
tion of a closterovirus associated with apple mealybug- 27(6):186191
transmitted little cherry disease in North America. Ward A (1966) Mealybugs (Hemiptera: pesudococcidae).
Phytopathology 91:268273 Acta Entomol Sinica 28:444446
Gilliatt FC (1935) A mealybug, Phenacoccus aceris Williams DJ (2004) Mealybugs of southern Asia. The
Signoret, a new apple pest in Nova Scotia. Can Natural History Museum/Southdene SDN. BHD.,
Entomol 67:161164 London/Kuala Lumpur, 896 p
Fruit Crops: Pears
30
M. Mani
Morimoto R (1976) A study of the biological control of Stokwe NF, Malan AP (2010) Potential use of entomo-
Comstock mealybug, Pseudococcus comstocki pathogenic nematodes for biological control of mealy-
Kuwana. Rev Plant Prot Res 9:7289 bugs on apples and pears. SA Fruit J 9(3):3839, 42
Myburg AC, Rust DJ, Stubbings D (1975) Mealybugs on Swart PL (1977) Mealybugs: more economic manage-
apples and pears. Deciduous Fruit Grower 25(7): ment on apples and pears. Deciduous Fruit Grower
176179 27(6):186191
Park DS, Leem YJ, Hahn KW, Suh SJ, Hong KJ, Oh HW Wakgari WM, Giliomee JH (2004) Mealybugs and their
(2010) Molecular identication of mealybugs parasitoids in apple and pear orchards in the Western
(Hemiptera: Pseudococcidae) found on Korean pears. Cape Province, South Africa. Afr Plant Prot
J Econ Entomol 103(1):2533 10(1):711
Seo HH, Lee JY, Jung HW (2010) Fruit appearance Warner G (2000) Mealybug shows up in soft fruit. From
improvement by using lter-attached paper bags in grapes, to pears, and now soft fruits, grape mealybug
Niitaka pears. Hortic Environ Biotechnol continues its advance. Good Fruit Grower 51(6):21
51(2):7377 Williams DJ (2004) Mealybugs of southern Asia. The
Natural History Museum/Southdene SDN. BHD,
London/Kuala Lumpur, 896 p
Fruit Crops: Plum
31
M. Mani
31.1 Species
chlorpyrifos-methyl, diazinon, dimethoate, mixture Uzbekistan laboratory, reared locally, and dis-
of dimethoate and methidathion, fenamiphos, imi- tributed in the Odessa region. This parasitoid had
dacloprid, methidathion, omethoate, oxydemeton- been already observed in Odessa, but it multi-
methyl, parathion, profenofos, and prothiofos. plied slowly, and regular releases were necessary
Postharvest treatments as high-volume sprays during the period of appearance of the second
(handguns) signicantly reduced insect popula- instar nymphs of each generation. Mass releases
tions. However, survival rates under bark as well of parasites had begun in 1977, and the effective-
as on plum roots made it necessary to apply com- ness reached 76.896.8 % in 1978. No releases or
plementary sprays in October and in December other control measures were undertaken in 1979,
January, both periods corresponding to extensive and infestation declined rapidly, parasitism being
migratory ows to the bunches and fruits. In this 98 %. Subsequent observations showed that
context, the control of P. afnis with trunk and soil Pseudaphycus readily became established in the
applications of chlorpyrifos proved necessary Odessa area and provided sufcient control for
(Gonzalez et al. 1996). Application of sprays of articial rearing and release to be discontinued.
diazinon, methidathion, and profenofos after the Since the outbreak appeared to have been due to
fruit harvest against Pseudococcus viburni (P. af- the importation of infested pomegranates, quar-
nis) was evaluated on table grapes and plums in antine measures were taken to ensure that such
Chile. These treatments considerably reduced pest imports were free from the mealybug
population levels. Nymphal mortality was greater (Romanchenko and Belskaya 1981).
than mortality of mature females (Gonzalez et al.
1995). In Chile, a new approach to minimize risks 31.3.2.3 Azerbaijan SSR, USSR
is suggested through control programs against P. In Apsheronsk Peninsula in the Azerbaijan SSR,
viburni in plums starting at the postharvest season, USSR, Phenacoccus mespili Sign was shown to
followed in the next early spring season with the be a pest of peach, apricot, quince, cherry plum
chitin inhibitor buprofezin. The use of neonicoti- [Prunus divaricata], cherry, bird cherry, apple,
noid insecticides is also under development to pear, and ash [Fraxinus]. The pest had two com-
include acetamiprid, imidacloprid, thiacloprid, plete generations and a partial one per year. The
and thiamethoxam (Gonzalez and Volosky 2004). rst generation developed in about 65 days and
The efcacy of spring and postharvest treatments the second in about 45.2 days. The most impor-
of insecticides (chlorpyrifos, ethoprophos, and tant natural enemy was the encyrtid Pseudaphycus
carbofuran) against P. viburni in plum orchards phenacocci Yasnosh that parasitized about 73.8
(cv. Larry Anne) in Chile is also discussed % of the pest population in late August and
(Gonzalez et al. 2001). September. Other parasitoids recorded were
Aphycus hadzibejliae Trjapitzin and Allotropa
mecrida (Walker), while the predators were
31.3.2 Biological Control Chilocorus bipustulatus L., Chrysoperla carnea
(Stephens), and Leucopis alticeps Czerny
31.3.2.1 California (Ibadova 1985).
In California, exotic parasitoids Allotropa burrelli
Mues., A. convexifrons Mues., and Pseudaphycus 31.3.2.4 Chile
malinus Gah were found to be successfully estab- Pseudococcus viburni (Ps. afnis) has become an
lished on Ps. comstocki (Meyerdirk and Newell economically important pest of Japanese plums
1979). in Chile. At harvest, a higher number of fruits are
infested inside the pedicel cavity. Mid-to-late
31.3.2.2 Crimea (USSR) season cultivars are often infested, and oviposit-
In the Odessa region of the Crimea (USSR), for ing females occur on fruits from mid-January.
control of the Comstock mealybug Ps. comstocki Postharvest treatments with chlorpyrifos,
on plum, Pseudaphycus was introduced from the methidathion, and a mixture of dimethoate and
31 Fruit Crops: Plum 309
chlorpyrifos are recommended for control of the afnis (Maskell) (Homoptera: Pseudococcidae), with
diazinon, methidathion and profenofos. Agric Tecnica
mealybugs (Gonzalez 1991).
(Santiago) 55(2):9598
Gonzalez RH, Curkovic ST, Barria PG (1996) Evaluation
of the efcacy of insecticides against mealybugs in
References plums and grapes (Homoptera: Pseudococcidae).
[Spanish]. Rev Frutic 17(2):4557
Gonzalez RH, Jorge Poblete G, Gerardo Barria P (2001)
Curkovic ST, Barria PG, Gonzalez RH (1995) Preliminary
The tree fruit mealybug in Chile, Pseudococcus
observations on insects and mites on grapes, pears,
viburni (Signoret) (Homoptera: Pseudococcidae).
plums and persimmons detected with corrugated band
Revista Fruticola 22(1):1726
traps [Spanish]. Acta Entomol Chilena 19:143154
Ibadova SI (1985) The apple mealybug Phenacoccus mes-
Godfrey LD, Pickel C (1998) Seasonal dynamics and
pili Sign and its natural enemies in the Apsheronskiy
management schemes for a subterranean mealybug,
Peninsula [Russian]. Izv Akad Nauk Azerbaidzhanskoi
Rhizoecus kondonis Kuwana, pest of alfalfa. Southwest
SSR Biol Nauk 1:3236
Entomol 23(4):343350
Meyerdirk DE, Newell IM (1979) Importation, coloniza-
Gonzalez RH (1991) Mealybugs (Homoptera:
tion, and establishment of natural enemies on the
Pseudococcidae), a new pest of Japanese plums in
Comstock mealybug in California. J Econ Entomol
Chile [Spanish]. Rev Frutic 12(1):37
72(1):7073
Gonzalez RH, Volosky FC (2004) Mealybugs and fruit
Richmond JE, Cowley JM (1998) Pineapple mealybug in
moth: quarantine problems affecting fresh fruit exports
New Zealand? Orchardist 71(7):71
[Spanish]. Rev Frutic 25(2):4162
Romanchenko AA, Belskaya NM (1981) The Comstock
Gonzalez RH, Curkovic ST, Barria PG (1995) Postharvest
mealybug in the Odess region [Russian]. Zashchita
control of the grape and plum mealybug Pseudococcus
Rastenii 4:41
Fruit Crops: Peaches
32
M. Mani
Comstock mealybug, Pseudococcus comstocki the second in about 45.2 days. The most impor-
(Kuwana), has been reported on peaches in tant natural enemy was the encyrtid Pseudaphycus
Illinois. Egg masses are commonly present on phenacocci Yasnosh which parasitized about
fruits, limbs, and bases of new shoots. 73.8 % of the pest population in late August and
Phenacoccus mespili Sign. is known to be a pest September. Other parasitoids recorded were
of peaches in the Azerbaijan SSR, USSR (Ibadova Aphycus hadzibejliae Trjapitzin and Allotropa
1985).In Apsheronskiy Peninsula in the mecrida (Walker), while the predators were
Azerbaijan SSR, USSR, the pest had two com- Chilocorus bipustulatus L., Chrysoperla carnea
plete generations and a partial one per year. The (Stephens) and Leucopis alticeps Czerny
rst generation developed in about 65 days and (Ibadova 1985).
Reference
Ibadova SI (1985) The apple mealybug Phenacoccus mes-
M. Mani (*) pili Sign., and its natural enemies in the Apsheronskiy
Indian Institute of Horticultural Research, Peninsula [Russian]. Izvestiya Akademii Nauk
Bangalore 560089, India Azerbaidzhanskoi SSR Biologicheskikh Nauk
e-mail: mmani1949@yahoo.co.in 1:3236
distributed more or less evenly on all buds. Neonicotinoids were the most toxic to the Pl.
Over the whole orchard, the frequency distribu- kraunhiae on persimmon, followed by organo-
tion of immature individuals appeared to cor- phosphates, while the synthetic pyrethroids were
respond to a concentrated distribution. Within a less effective in Japan (Morishita 2006).
tree, the frequency distribution of the number A mean LT 99 of Ps. longispinus at 44 C was
of individuals per bud varied with density; 74.2 min, which decreased to 15.1 min at 54
when it was low, it corresponded to Poissons C. Hot water immersion appeared to be a poten-
curve, but when it was high (more than about tially useful disinfestation method. The mortality
0.5/bud), it could be tted to the concentrated response of Ps. longispinus on persimmons to hot
type. The same trend was observed in both spe- water immersion treatments between 44 and 54
cies (Ueno 1971). C was examined. The calyx of the persimmon
Ferrisia gilli, Gills mealybug, is a newly was found to offer thermal protection for
described species of mealybug that is spreading P. Longispinus, resulting in lower insect mortal-
throughout California, infesting many stone ity under the calyx compared to that on the out-
fruits and also persimmons (Gullan et al. 2003). side of the fruit (Lester et al. 1995). Koide et al.
Parasitoids that include wasps in the genera (2009) predicted the hatch timing of the mealy-
Pseudaphycus, Chrysoplatycerus, and Anagyrus bug Pl. kraunhiae in persimmon orchards using
have been shown to effectively reduce Gills the effective accumulated temperature calcula-
mealybug populations on persimmon crop sys- tion simulation of the JPP-NET in Aichi.
tems where pyrethrin-based insecticide use is Hot air treatment of P. longispinus on persim-
very limited. mons achieved 99 % mortality of the mealybug
with 12.4 h at 44 C, which reduced to 4.5 h at 47
C and 3.8 h at 50 C (Dentener et al. 1996).
33.2 Management Treatment at a 47 C-persimmon esh tempera-
ture for up to 3 h after a 2-h warm-up period, fol-
Application of chlorpyrifos (5 %) and diazinon (4 lowed by immediate cold storage, has the
%) applied around the trunk gave excellent results benecial effect of delaying the onset of chilling
against ants. All the treatments greatly reduced injury in persimmons while causing only slight
the population of mealybug P. citri (Izhar 1999). internal and external damage to the fruit (Woolf
There are three generations of Japanese mealy- et al. 1997). Therefore, a combined heat-to-cold
bug, Planococcus kraunhiae, on persimmons storage treatment may be effective for disinfesta-
each year. Chemicals are to be applied usually in tions of P. longispinus on persimmons. An esti-
June and August when rst instar nymphs, the mated treatment time of 3.3 h (including a 2-h
most susceptible to insecticides, appear. The den- warm-up period) at 44 C, followed by a 40-day
sity of Japanese mealybug, Planococcus cold storage at 0 C, was needed to achieve 99 %
kraunhiae, on Japanese persimmon fruit was mortality (Dentener et al. 1997).
higher in plots frequently treated with cyperme- Cryptolaemus montrouzieri was introduced
thrin than that in the untreated plot. The number into Japan for the control of Planococcus
of mealybugs found on Fuyu, a non-astringent kraunhiae on persimmon (Ishi 1940).
cultivar, was higher than that on Hiratanenashi,
an astringent cultivar. The following additional
control measures should be taken in heavily References
infested orchards: (1) eliminating overwintering
nymphs by scraping away the tree bark, (2) Charles JG (1993) A survey of mealybugs and their natu-
ral enemies in horticultural crops in North Island, New
spraying the tree with petroleum oil in winter,
Zealand, with implications for biological control.
and (3) applying pesticides from late April to Biocontrol Sci Technol 3(4):405418
early May when overwintering nymphs move to Curkovic ST, Barria PG, Gonzalez RR (1995) Preliminary
the top of shoots (Morishita 2005a, b). observations on insects and mites on grapes, pears,
33 Fruit Crops: Persimmon 315
plums and persimmons detected with corrugated band Morishita M (2005b) Resurgence of Japanese mealybug,
traps [Spanish]. Acta Entomol Chilena 19:143154 Planococcus kraunhiae (Kuwana), in persimmon
Dentener PR, Alexander SM, Lester PJ, Petry RJ, induced by a synthetic pyrethroid cypermethrin
Maindonald JH, McDonald RM (1996) Hot air treat- [Japanese]. Ann Rep Kansai Plant Prot Soc
ment for disinfestation of lightbrown apple moth and 47:125126
longtailed mealy bug on persimmons. Postharv Biol Morishita M (2006) Susceptibility of the mealybug,
Technol 8:143152 Planococcus kraunhiae (Kuwana) (Thysanoptera:
Dentener PR, Bennett KV, Hoy LE, Lewthwaite SE, Thripidae), to insecticides evaluated by the petri dish-
Lester PJ, Maindonald JH, Connolly PG (1997) spraying tower method [Japanese]. Jpn J Appl
Postharvest disinfestation of light brown apple moth Entomol Zool 50(3):211216
and longtailed mealybug on persimmons using heat Prestidge RA, Holland PT, Clarke AD, Malcolm CP
and cold. Postharv Biol Technol 12:255264 (1989) Pesticides for use close to and during harvest of
Dunkelblum E, Zada A, Gross S, Fraistat P, Mendel Z persimmons. In: Popay AJ (ed) Proceedings of the
(2002) Sex pheromone and analogs of the citrus 42nd New Zealand Weed and Pest Control Conference,
mealybug, Planococcus citri: synthesis and biological New Plymouth, New Zealand, pp 195199
activity. Bull OILB/SROP 25(9):213220 Steven D, Sale PR (1985) Insect control trials on persim-
Gullan PJ, Downie DA, Steffan SA (2003) A new pest mons. In: Hartley MJ (ed) Proceedings of the 38th
species of the mealybug genus Ferrisia Fullaway New Zealand Weed and Pest Control Conference,
(Hemiptera: Pseudococcidae) from the United States. Rotorua, New Zealand, pp 203206
Ann Entomol Soc Am 96:723737 Tranfaglia A (19721973) Studies on Homoptera
Ishi T (1940) The problem of biological control in Japan. Coccoidea I on the discovery in Campania of
Proc 6th Pac Sci Congr 4:365367 Pseudococcus obscurus Essig, a species new to the
Izhar J (1999) Preliminary experiment to control ants as a Italian fauna [Italian]. Bollettino del Laboratorio di
transferring factor of millibug (Pseudococcus citricu- Entomologia Agraria Filippo Silvestri Portici
lus, Planococcus citri) in persimmon [Hebrew]. Alon 30:294299
Hanotea 53(12):490492 Tsutsumi T (1997) Effect of mospilan on the control of
Koide T, Suzuki H, Kuwayama S, Yamaguchi K (2009) Japanes mealybug on persimmon tree (Japanese).
Prediction of the hatch timing of the mealy bug Noyaku Jidai 174:2729
Planococcus kraunhiae (Hemiptera: Pseudococcidae) Ueno H (1971) Studies on the scale insects that attack the
in persimmon orchards using the effective accumu- Japanese persimmon Diospyros kaki
lated temperature calculation simulation of the JPP- L. II. Interrelationships between overwintered larvae
NET in Aichi [Japanese]. Ann Rep Kansai Plant Prot of the Japanese wisteria cottony mealybug,
Soc 51:115116 Planococcus kraunhiae Kuwana, and the elongate cot-
Lester PJ, Dentener PR, Petry RJ, Alexander SM (1995) tony scale, Phenacoccus pergandei Cockerell, in the
Hot-water immersion for disinfestation of lightbrown orchard [Japanese]. Jpn J Appl Entomol Zool
apple moth (Epiphyas postvittana) and longtailed 15(4):211214
mealy bug (Pseudococcus longispinus) on persim- Woolf AB, Ball S, Spooner KJ, Lay Yee M, Ferguson IB,
mons. Postharv Biol Technol 6(3/4):349356 Watkins CB, Gunson AF, Forbes SK (1997) Reduction
Morishita M (2005a) Effect of bark-scraping, dormant of chilling injury in the sweet persimmon Fuyu dur-
spray of petroleum oil and applying pesticide in late ing storage by dry air heat treatments. Postharv Biol
spring on density of Japanese mealybug, Planococcus Technol 11:155164
kraunhiae (Kuwana), in persimmon [Japanese]. Ann
Rep Kansai Plant Prot Soc 47:123124
Fruit Crops: Passion Fruit
34
M. Mani
34.1 Species
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Pseudococcus maritumus (Ehrhorn) is a problem The damage caused to apricots is due to excretion
on apricot in the USA (Anonymous 1980). of honeydew by Ps. maritumus. Colonies are
Phenacoccus mespili (Sign.) is known to be a formed in the depression around the stem end of
pest of apricot in the Azerbaijan SSR, USSR the fruits, and honeydew produced run over sides
(Ibadova 1985). Phenacoccus aceris (Signoret) of apricots. The black smut fungus that grows in
is also known to attack all deciduous fruit and nut the honeydew gives the fruit an unsightly appear-
trees, including apricots in Nearctic and ance. In addition, the honeydew gives the fruit a
Palaearctic regions (Ben-Dov 1994). Ferrisia vir- reddish tint. As apricots are picked relatively
gata (Cockerell) is also known to attack apricots ripe, it is not possible to remove the honeydew by
in Egypt. the normal washing procedure. The fruit is not
suitable for fresh shipment, and processors of
unpeeled halves consider the contaminated fruit
as culls.
nymphs are pink to purple with waxy laments As dormant spray, horticultural mineral oil is rec-
giving them a whitish cast. Adult females resem- ommended at 12 gal/100 gal water (48
ble nymphs and are about 0.19 in. long and quite gal/a) + diazinon (Diazinon AG500) at 1 pint/100
mobile. Later in the season, mealybugs may pro- gal water (1.53 pints/a) for control of mealybug.
duce copious amounts of honeydew. They reach A prebloom spray application of insecticides
maturity by MayJune. Receptive females release diazinon (50 W) at 1 lb/100 gal water (4 lb/a) and
a pheromone to attract males. Adult males appear phosmet (70 W) at 0.751 lb/100 gal water (4.25
rst, mate with last instar nymphs or adult lb/a) is given before leaves begin to curl, and
females and die, and females deposit eggs in the before petal fall. Diazinon (50 W) at 1 lb/100 gal
cracks of the bark. The eggs hatch and crawlers water (4 lb/a) and imidacloprid (1.6 F) were rec-
of the second generation move to both foliage ommended at 2 oz/100 gal water (48 oz/a) is
and fruit during June and early July. It is at this recommended as petal fall spray for the control
time that the mealybugs colonize around the stem of mealybugs (Madsen and Mcnelly 1960).
end of the fruit. Apricots are usually harvested in
July, and after the fruit is picked, the mature,
mated females migrate to sheltered areas, lay References
eggs, and die in the egg sac. The eggs hatch in
September, but the crawlers remain within the Anonymous (1980) Pseudococcus maritimus [Distribution
map]. Distribution Maps of Plant Pests 1980. Map 404
old egg mass until the following spring (Madsen
Ben-Dov Y (1994) A systematic catalogue of the mealy-
and McNelly 1959). bugs of the world (Insecta: Homoptera: Coccoidea:
Pseudococcidae and Putoidae) with data on geograph-
ical distribution, host plants, biology, and economic
importance. Intercept Limited, Andover, 686 p
35.4 Management Ibadova SI (1985) The apple mealybug Phenacoccus mes-
pili (Sign.) and its natural enemies in the Apsheronskiy
Treatments timed to the spring emergence of Peninsula [Russian]. Izvestiya Akademii Nauk
crawlers were effective and were preferred to fall Azerbaidzhanskoi SSR. Biologicheskikh Nauk
1:3236
or winter sprays. Diazinon was found to be effec-
Madsen HF, McNelly LB (1959) Mealybug on apricot.
tive against the mealybug on apricots. Weekly Calif Agric 13:1314
sprays of horticultural oil, neem oil, and use of Madsen HF, McNelly LB (1960) Control of the grape
insecticidal soap work well against mealybugs. mealybug on apricots. J Econ Entomol 53(3):354357
Fruit Crops: Pistachio and Almond
36
M. Mani
Gills mealybug Ferrisia gilli Gullan is a primary Mealybug feeding results in the production of
pest of pistachio in California. Mealybug infesta- large amounts of honeydew that acts as a sub-
tion causes a decrease in nut quality because of strate for black sooty mold. Stems, leaves, and
increased shell staining and possibly smaller ker- clusters in trees are often covered in honeydew
nel size. Mealybug populations are at their high- and sooty mold. Thick layers of sooty mold on
est at the time of harvest. Ferrisia gilli is a leaf surfaces reduce photosynthesis. Mealybugs
relatively large mealybug that feeds by sucking have a great afnity for feeding within the pis-
plant juices of almond in California. Mealybug tachio cluster. They use piercingsucking
control is achieved through a dormant or June mouthparts to suck out plant juices, extracting
application of buprofezin, which is highly effec- carbohydrates and other nutrients intended for
tive against immature stages and can reduce nut development. This causes a decrease in nut
mealybug populations in a manner that is rela- quality because of increased shell staining and
tively safe to predators and parasites. possibly smaller kernel size. During the late
Alternatively, chlorpyrifos provided excellent spring through harvest, mealybug is particu-
control when sprayed in the dormant season and larly found feeding within the cluster where
would likely do the same in season. Parasitoids they cause losses in quality and possibly yields.
include wasps in the genera Pseudaphycus, Harvesting is also affected when severe hull
Chrysoplatycerus and Anagyrus. These parasit- damage causes nuts to dry up and shrivel on
oid species have been shown to effectively reduce the tree.
Gills mealybug populations where pyrethrin-
based insecticide use is very limited.
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Monitoring of mealybugs on the trees is an Several species of predators and parasitoids can
important step to make treatment decisions. At suppress Gills mealybug densities. Predators
budbreak, search for mealybugs is to be done at include green lacewings and a small brown coc-
the bases of new buds on trees known to be previ- cinellid (ladybird) beetle whose larva mimics the
ously infested. Treatment decisions are to be appearance of a mealybug. Parasitoids include
made by determining the number of adult female wasps in the genera Pseudaphycus,
mealybugs per cluster in late May. An average of Chrysoplatycerus, and Anagyrus. These parasit-
three mealybugs per cluster in May is sufcient oid species have been shown to effectively reduce
to cause a 15 % reduction in the value of the crop Gills mealybug populations where pyrethrin-
at harvest. It is advised to look for mealybug based insecticide use is very limited.
infestations in fall after harvest, and mark areas
in the orchard where they occur so that their pop-
ulations can be monitored the following spring. If
adult females are found in clusters in May, a
treatment aimed at crawler emergence may be
warranted.
The best time to nd new mealybug infesta-
tions is the period from early fall through mid-
winter when populations are at their highest.
Before trees become dormant, it is advised to
look for sooty mold on leaves and for mealybugs
within the clusters. Once the leaves have fallen,
look for white aggregations of mealybugs on the
trunks and undersides of main scaffolds. If
mealybugs are found, mark and follow up on
these locations the following spring. Parasitized mealybug
The most effective timing for insecticides is
when most mealybugs are in the crawler stage of
the rst generation, which for the lower San
Joaquin Valley is around early to mid-June. Be
sure to monitor clusters to determine crawler
emergence. Applications later in the season are
more variable in effectiveness. Postharvest treat-
ments are not recommended because this is when
biological control is most active, no damage
occurs to the crop in winter, and there is already
high winter mealybug mortality. The insecticide
buprofezin 34.5 oz is very effective when used
while mealybugs are in the crawler stage of the
rst in-season generation. Acetamiprid 2.3
4.1 oz is effective against second-generation
crawlers in mid-to-late July. Predatory larva on the mealybug
324 M. Mani
secretions that cover their bodies. The California control is achieved through a dormant or June
gray ant (eld ant) also interacts heavily with F. application of buprofezin, which is highly effec-
gilli. Field ants are attracted to mealybugs and tive against immature stages and can reduce
were often found in close association with them. mealybug populations in a manner that is rela-
It is likely that predation on the crawler stages tively safe to predators and parasites.
that appeared in mid-June could explain the Alternatively, chlorpyrifos provided excellent
abrupt disappearance of the mealybugs for the control when sprayed in the dormant season and
remainder of the season, especially since there would likely do the same during in-season.
were lots of eld ants, no insecticides were used, Dormant treatments, however, would be pre-
and there were no mealybug mummies left ferred since they should be relatively safe to para-
behind that would indicate populations were sitoids due to their state of dormancy inside of
reduced through parasitism. mealybug mummies.
Trees treated with chlorpyriphos were the only Gullan PJ, Downie DA, Steffan SA (2003) A new pest
species of the mealybug genus Ferrisia Fullaway
trees to have signicant reductions compared to
(Hemiptera: Pseudococcidae) from the United States.
the untreated control. During April and May, Ann Entomol Soc Am 96:723737
once overwintering mealybugs had molted, it Shaheen G, Zahoor B, Ahmad AS (2014) Management of
resulted in an excellent control of the pest by almond mealybug, Drosicha dalbergiae (Stebbing) by
chemical intervention. Ann Plant Prot Sci
buprofezin, followed by chlorpyriphos. Mealybug
22(1):2226
Fruit Crops: Strawberry
37
M. Mani
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Mealybugs have been reported as serious pests in and Planococcoides robustus sp.n. (Ezzat and
North America, South America, Canada, Mexico, McConnell) in Karnataka in 1976 (Puttarudraiah
USSR, France, South Africa, Australia, Italy, and Murthy 1976). Prior to 1980, occasional
New Zealand, Chile, Middle East countries, etc. losses occurred as a result of localized infesta-
(Table 38.1). Economic losses resulting from tion, and usually disappeared in the following
mealybug infestations on grapes have dramati- year. But in the early 1980s, economic losses on
cally increased in India. As many as seven spe- grapes in Andhra Pradesh, Maharashtra,
cies are known to attack grapevine in India (Mani Karnataka and to some extent in Tamil Nadu led
et al. 2008). Mealybugs are considered to be the to the rediscovery of pink hibiscus mealybug,
most important pests of grapevine in India par- which was also reported on a wide range of host
ticularly in Maharashtra, Andhra Pradesh, Tamil plants in peninsular India. From the mid-1980s
Nadu and Karnataka. onwards, mealybugs have become persistent
Historical perview on mealybugs in India pests in peninsular India (Satyanarayana 1981;
that the pink hibiscus mealybug Maoconellicoccus Mani 1986; Reddy and Narayana 1986; Azam
hirsutus (Green) as Phenacoccus hirsutus 1983; Srinivasan 1987). Extensive use of insecti-
(Green) was rst reported on grapes in 1919 cides in vineyards might have resulted in the out-
(Fletcher 1919, 1923), spherical mealybug break of mealybugs in the late 1980s (Manjunath
Nipaecoccus viridis (Newstead) as Pseudococcus 1985). Grape production is often adversely
corymbatus (Green) in 1932 (Fletcher 1932) and affected due to the mealybugs, with the extent of
also as Pseudococcus lamentosus (Cockerell) in damage being as much as 90 % in extreme cases.
Punjab in 1946 (Anonymous 1946), striped Apparently, it could be due to the disruption of
mealybug Ferrisia virgata (Cockerell) in Tamil natural enemies of mealybugs by broad-spectrum
Nadu in 1958 (Raman 1958), N. viridis in 1965 insecticides. In fact, mealybug infestation
(Subba Rao et al. 1965) and Pseudococcus sp. in increased with the increased use of insecticides,
Andhra Pradesh in 1974 (Tejkumar et al. 1977) particularly organophosphates.
Table 38.1 Mealybug species recorded on grapevine in different regions of the world
Species Region Reference
Asterococcus muratae (Kuw.) USA Lambdin (1983)
Dysmicoccus brevipes (Ckll.) Brazil Daane et al. (2012)
India Mani (1986)
Ferrisia malvastra (McDaniel) S. Africa Walton and Pringle (2004); Iordanou (1974)
Argentina Tryapitzyn and Tryapitzyn (1999)
Ferrisia gilli (Gullan) USA Gullan et al. (2003)
Geococcus coffeae (Green) Ben-Dov (1994)
Heliococcus bohemicus (ulc.) Hungary Jakab and Szendrey (1989)
Italy Camporese (1994)
France, Germany Sforza et al. (2003)
Switzerland Kozar et al. (1994)
Macoonellicoccus hirsutus (Green) India Reddy and Narayana (1986)
Nipaecoccus viridis (Newstead) India Mani (1986)
Peliococcus turanicus (Kritshenko) Palaearctic region Ben-Dov (1994)
Phenacoccus aceris (Sign.) Italy Jablonowski (1917)
Phenacoccus hystrix (Baer) Germany Wunn (1928); Zillig and Neimeyer (1929)
Phenacoccus madeirensis (Green) Yemen Marotta et al. (2001)
Phenacoccus solani (Ferris) S. Africa Walton and Pringle (2004); Iordanou (1974)
Planococcus citri (Risso) Egypt Bodenheimer (1944)
France Bonnemaison (1962)
Hungary Anonymous (1917)
Italy Jablonowski (1917)
Israel Avidov and Swirski (1950)
Spain Ruiz Castro (1938); Cabaleiro and Segura (1997)
S. Africa Joubert (1943)
Turkey Aykac and Erguder (1972)
UK Brotherston (1914)
USA
California Golino et al. (2002)
USSR Pintz (1932); Chochiya (1941); Rozanov and
Loseva (1963); Niyazov (1969); Kurdyukov and
Alan (1973)
Brazil Morandi Filho et al. (2007); Cabaleiro and
Segura (1997)
Chile Gonzalez (2003); Artigas (1994)
Australia CSIRO (2001)
Planococcus cus (Sign.) France Panis and Trevillot (1975)
Italy Transfaglia (1976); Forte et al. (2008)
S. Africa Whitehead (1961); Walton and Pringle (2004b)
USSR Dantsig (1977)
Yemen Marotta et al. (2001)
Iran
Iraq, Israel, Lebanon,
Libya, Egypt
Tunisia Mahfoudhi and Dhouibi (2009)
Turkey Kaydan and Klncer (2005)
(continued)
38 Fruit Crops: Grapevine 331
In the mid-1990s, the spherical mealybug leaves and shoot tips. As the mealybugs feed,
Nipaecoccus viridis (Newstead) was also reported they excrete carbohydrate-rich honeydew that
to cause occasional losses in some vineyards in also serves as a substrate for the growth of sooty
South India. A localized infestation of mould on leaves, shoots and bunches. Sooty
Xenococcus annandalei (Silvestri) in 1996 was mould inhibits photosynthesis and affects the
also reported in North Bangalore (Rajagopal growth and development of vine. Second, it
et al. 1997) and Planococcus minor (Maskell) adversely affects the fermentation process and
(Planococcus pacicus (Cox) in Punjab (Batra subsequently taints the wine. The damage pro-
et al. 1987). But in mid-2000, citrus mealybug duced by mealybugs is due to the presence of
Planococcus citri (Risso) was reported to cause one or more of the following: the cottony ovi-
severe losses in Maharashtra and Karnataka par- sac, eggs, nymphs, adults, honeydew or sooty
allel to M. hirsutus (Mani and Kulkarni 2007). mould. Honeydew often drips onto the fruit
Due to awareness on the use of harmful broad- from the mealybugs feeding on the foliage
spectrum insecticides by farmers, increased use above the clusters. Honeydew is colourless and
of selective chemicals and biopesticides, grape syrupy when rst exuded; it later becomes
mealybug populations decreased noticeably in darker because of the sooty mould. Grape ber-
late 2000 in India. Although individual vineyards ries in an infested bunch do not develop nor-
suffered losses due to mealybugs, the problem mally and are shrivelled. Bunches having sooty
became considerably less severe, and in many mould-coated berries will be unsightly, thereby
cases treatments were reduced or eliminated. losing its market value due to cosmetic damage
Still, individual vineyards suffer from mealy- to the grape clusters; they are poor in quality
bugs, which require treatments. and unt for human consumption. The grape
mealybug alone caused 50100 % yield losses
in the eld (Azam 1983). The pest attack weak-
38.1 Damage ens the grown-up vines. The mere presence of
mealybug colonies and sooty mould causes cos-
Mealybugs are phloem feeders that use long, metic damage to grape cluster. In case of severe
slender mouthparts to suck the sap from the mealybug infestation, young vines often die.
trunk, cordons, buds, spurs, aerial roots, leaves, Difference in the amount of damage caused by
shoots, nodes, ower panicles and bunches. each species is often related to population size,
Infestation of the growing point, especially with preferred feeding locations and temperature
the pink mealybug, results in malformation of tolerances.
Honeydew can be dissolved by light rain and roots; and the affected vines show reduced
will dry in warm temperatures; however, when vigour, shortening of fruit-bearing canes and
infestations are very severe, it can accumulate to reduction in size of fruit bunches and yield.
form a hard, wax-like layer that covers the Like most other grape pests, grape mealybugs
infested plant material and results in defoliation, prefer vigorous vines. Thus, vines most likely to
and repeated annual infestations result in vine be infested are outside rows, since these are nor-
death. Fruit clusters in direct contact with spurs mally the most vigorous. Weak vines may har-
or trunk are more likely to be damaged. bour mealybugs, but heavy populations are
Generally, the table grapes suffer very heavily normally found mainly on fairly vigorous vines.
in comparison with raisin and wine grapes. The In general, the mealybug infestations are con-
table grape vineyards are usually more easily ned to few vines, while others are clean. But
infested because of the greater use of pesticides when there is outbreak, all the vines are likely to
to ensure clean fruits. This sometimes interferes be infested. Many a time, the infestation is local-
with natural control factors. Area under wine ized. In a given area, one vineyard may be heav-
grapes has started increasing recently in India. ily infested while many others may be completely
The mealybug also poses a serious threat to free from mealybugs under the same conditions.
wine grapes. It is very difcult to process the Grape bunches that touch old wood have signi-
fruits for raisin and wine if the bunches are cantly higher damage and mealybug densities.
heavily infested with mealybugs. The root The majority of mealybugs are always found in
mealybugs Xenococcus annandalei and protected locations (under the bark of the trunk,
Planococcoides robustus in India also cause spurs or canes), indicating the need for chemical
damage occasionally by sucking the sap from or biological controls that can penetrate these
334 M. Mani and U. Amala
refugia (Chris and Kent 2001). Mealybugs are mealybug population in JulyAugust (Mani
also known to transmit Grapevine-leafroll- 1986; Balikai 1999b).
associated virus (GLRaV) on grapevine in many The seasonal incidence of mealybug was
countries. Recently, the diseases have been found expressed in terms of standard weeks. A popula-
damaging wine grapes in peninsular India. The tion of 25.0 colonies per vine was observed dur-
vine mealybug as a vector of GLRaV is yet to be ing the 14th standard week, and thereafter it
established in India. declined and reached to a minimum of 7.4 colo-
nies during the 22nd standard week due to April
pruning effect. From the 24th standard week,
38.2 Seasonal Development again it started increasing and reached to a peak
during the 36th standard week (14.5 colonies/
In India, the mealybug occurs on the grapevine vine). From the 39th standard week, it again
throughout the year. Seasonal development of started declining and reached to a minimum of
mealybugs depends on the phenology of the crop. 5.0 colonies during the 44th standard week due to
Development of mealybug population can be September pruning effect and once again started
related to vine development. After pruning in increasing steadily in the fruiting season and
SeptemberOctober (fruit pruning), the mealy- reached a peak of 32.4 colonies per vine during
bugs remain low on the trunk, cordons and stem the 10th standard week (Katke 2008) (Fig. 38.2).
up to the rst fortnight of December. In general, A similar type of seasonal development of M.
the mealybug population starts increasing from hirsutus was observed on grapes in Andhra
mid-December onwards. During January, they Pradesh (Azam, 1983: Babu and Azam 1987) and
migrate from the trunk, cordons and shoots to also in Maharashtra (Anonymous 1992; Koli
ower panicles and then developing berries. It 2003). The development of other important
attains its peak population before the harvest of mealybugs like Planococcus citri and
bunches during MarchApril. The grapevine is Nipaecoccus viridis also follows a similar pattern
pruned usually in AprilMay (foundation prun- in south India.
ing) (Fig. 38.1). Mealybugs remain on the leaves, Fletcher (1919) reported that M. hirsutus had
stem and trunk from April to September. The ten generations per year in India. The number of
mealybug population is usually low from June to generations of the mealybug varies with the spe-
September coinciding with the rainy season. In cies, locality and climatic factors. There is also
the absence of rains, there is a sudden spurt in the variation in seasonal feeding, location and
500
Number of mealybugs / plant
450
400
350
300
250
200
150
100
50
0
Oct
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
movement on the vine among species and within 38.3 Varietal Susceptibility
species depending on regional temperatures and
vineyard management practices. There is no dia- Grape varieties that produce clusters close to the
pause, and slow development may occur during base of shoots so that the fruit touches the old
the winter months under south Indian conditions. wood are likely to have more heavily infested
Temperature is the driving force for mealybug clusters than varieties where clusters hang more
development (e.g., Chong et al. 2008), although freely. Early-maturing varieties are much less
development times and temperature thresholds likely to have serious fruit damage than the late-
differ among the species. Heavy sporadic rains maturing varieties. Cane-pruned varieties are a
and cool temperatures of less than 20 C result in little less likely to be infested seriously than spur-
temporary reduction in the mealybug population. pruned canes. Among commercial grape variet-
The pest population build-up coincides with high ies, none was found free from its attack. Seedless
temperature of 3040 C, low humidity (less than cultivars with tight lling of the clusters have
40 %) and berry development. more infestation than the seeded and loosely
The population is low in winter and rainy lled clusters. Raman (1958) observed more inci-
seasons and higher in summer months (Reddy dences of mealybugs in Pachadraksha (Bhokri)
and Narayana 1986; Babu and Azam 1987; and Thomspon in seedless ones but less incidence
Mani and Thontadarya 1987a, b; Manjunath in Black Prince, Phakdi, Anab-e-Shahi and
1985). There was a highly signicant positive Bangalore Blue. More or less incidence of mealy-
correlation of maximum and minimum tempera- bug in different varieties could also be a chance
ture and highly signicant negative correlation factor.
of morning and evening relative humidity and
non-signicant negative correlation of rainfall
with mealybug population in vineyards (Mani 38.4 Monitoring
and Thontadarya 1987a, b; Koli 2003). They
manage to survive under loose bark, feeding at The control strategy of this pest requires an effec-
bases of spurs, callus tissue at the site of girdles tive monitoring of the population dynamics,
in the off season. which is affected by various abiotic and biotic
336 M. Mani and U. Amala
parathion provided grape-mealybug control susceptible than the larger mealybugs. This is
(Frick 1952). Eventually, these materials became especially true for insecticides with a short resid-
less effective (Flaherty et al. 1982) and many ual period. Most research, therefore, has been
were ultimately banned from use. aimed at proper application timing and develop-
Many organophosphates are still effectively ing materials with better penetration into the
used (Gonzalez et al. 2001; Walton and Pringle mealybugs protected habitats. Dormant season
2001; Sazo et al. 2008). Newer materials, with or early spring application takes advantage of the
more novel modes of action, have also gained leaess vine, but mealybugs are in more pro-
popularity and include neonicotenoids, insect- tected locations. Applications with systemic
growth regulators, botanicals and biosynthesis materials near bloom are often used as the insec-
inhibitors (Daane et al. 2006; Sunitha et al. 2009; ticide moves out quickly to the leaves. After
Lo and Walker 2010). A major difference bloom, foliar materials should be applied beneath
between the older and new materials is coverage. the leaf canopy and aimed towards the grape
As mentioned, a portion of the mealybug popula- clusters and interior canes. In addition to the pos-
tion is often under the bark and, for some species, sibility of berry spotting, fresh market table
on the vine roots. Many of the older foliar sprays grapes possess a dull haze or dust on the skin,
do not effectively contact and kill mealybugs in termed bloom and the use of some insecticides
these more protected locations. The more novel can remove the bloom and lower the crop value.
materials have systemic properties, applied either Nevertheless, insecticides are the primary control
through the irrigation system or as a foliar. For tool for the mealybug control. Chemicals are to
organic or sustainable farming programmes, be applied through soil or can be sprayed to check
neem, light mineral oils, lime sulphur, citrus the mealybug populations.
products and fatty acid soaps have been used, but
these products have provided mixed results 38.5.3.1 Foliar Applications
(Srinivas et al. 2007). It is the most common method of applying insec-
Another historical difference is that the earlier ticides to control the mealybugs. Chemicals are
materials were often broad spectrum and killed effective if the sprays are applied when the mealy-
more than just the targeted mealybugs. Flaherty bugs are in the dispersive crawler stage and when
et al. (1982) stated that extensive use of DDT and the food plant affords the least shelter. Treatment
other synthetic insecticides to control leafhop- before the bud-burst stage and again after ower-
pers, etc. apparently disrupted the natural control ing reduces the mealybug population below the
of grape mealybugs. Other researchers have since economic threshold. Sprays of methidathion and
discussed the impact of broad-spectrum insecti- dimethoate against Pseudococcus maritimus
cides on mealybug natural enemies (e.g., Mani (Ehrh.) on grapevines (of the Thompson seedless
1986; Walton and Pringle 2001; Mgocheki and variety) in California resulted in 100 % control
Addison 2009). The cosmopolitan goal of man- (AliNiazee and Stafford 1972). The best control
aging vineyards with fewer broad-spectrum pes- of P. citri on grapevines was obtained with
ticides along with the development of resistance dimethoate, malathion or trichlormetaphos all at
to common pesticides (Flaherty et al. 1982; 0.2 % (Kurdyukov and Alan 1973). Three sprays
Charles et al. 1993) has fuelled the use of more of 0.075 % dichlorvos for controlling mealybugs
novel materials and research to improve mealy- (Pseudococcus sp.) on table grapes were applied.
bug controls. The rst was applied seven days before and the
For most materials, application timing is criti- second was 7 days after the beginning of harvest;
cal (Daane et al. 2012). Control measures are to the third spray 2 weeks later enhanced control
be taken at bud-burst stage, if any mealybugs are (Swart and Barnes 1975).
found during the previous harvesting. Exposed Currently, in North America, insecticide pro-
mealybugs are more easily killed than those grammes are based on the use of one or more of
under the bark, and the smaller stages are more the following insecticides: imidacloprid (a neo-
38 Fruit Crops: Grapevine 339
nicotinoid applied as a systemic near-bloom lowest mealybug population with increased cost-
time), buprofezin (an insect-growth regulator to-benet ratio (Biradar et al. 2006). Methomyl
applied as a foliar in late spring or early summer), @ 500800 g a.i/ha was found to be very effec-
acetamiprid (a foliar-applied neonicotinoid tive and gave high returns (Raguraman and
applied from late spring to harvest), clothianidin Premalatha 2006).
(a third-generation neonicotinoid applied as
either a foliar or systemic from late spring to har- 38.5.3.2 Dipping of Grape Bunches
vest), spirotetramat (a tetramic acid that acts as a Dipping of grape bunches for two minutes in any
lipid biosynthesis inhibitor and is applied from one of the insecticides, namely phosalone (2
late spring to early summer, or as post harvest) mL/L), monocrotophos (1.25 mL/L) or dichlor-
and chlorpyriphos (an organophosphate that is vos (mL/L) mixed with 25 g/L of sh-oil soap,
still used as a delayed dormant or post-harvest was highly effective in controlling mealybugs on
application) (Bentley et al. 2008). Prothiophos at bunches. Dipping in insecticide solution mixed
30 g/100 L or 1 mL/L afforded very effective with sh-oil rosin soap resulted in the scorching
control of the mealybugs throughout the season of berries at the blossom end due to the accumu-
(Prince and Fisher 1982). In the United States, lation of the mixture, but spraying was safe.
foliar application of buprofezin and chlorpyri- Though efcacy of insecticides was more by dip-
phos brought 82.7 and 85.0 % reduction in clus- ping than spraying with dichlorvos, both the
ter damage. Buprofezin is less expensive and methods of application were equally effective
provides excellent control. It is an insect-growth (Reddy and Narayana 1986).
regulator, most effective against smaller mealy-
bugs undergoing insect moults (Daane et al. 38.5.3.3 Soil Drenching
2008b). In India, dichlorvos was the most com- Chemigation (application of chemicals through
monly recommended chemical against mealy- irrigation) is an environmentally safe and the
bugs (FIP 1982). Foliar application of buprofezin most effective to control mealybugs. Imidacloprid,
at 1125 ml/ha reduced the nymphal and adult a systemic transluminar insecticide and also thio-
populations and bunch infestation of M. hirsutus methoxam (applied through the irrigation water
and increased the fruit yield (Muthukrishnan and taken by the vine roots), has been used in
et al. 2005). Buprofezin 25 SC @ 1125 ml/ha several countries and excellent control of mealy-
along with sh-oil rosin soap @ 3125 g/ha fur- bug has been obtained for a longer time.
ther improved the control of mealybugs on Imidacloprid provides greatest reduction of
grapes. Application of buprofezin does not affect 9093 % in cluster damage when applied through
the locally occurring natural enemies of vine drip irrigation (Daane et al. 2008b). In the drip-
mealybugs in India. Three sprays commencing irrigated vineyards, a four to six pretreatment
from the rst fortnight of January and subsequent irrigation prepares the soil; imidacloprid is then
sprays applied at ten days interval with dimetho- applied through irrigation system, and 6- to 8-h
ate 30 EC at 1.7 mL + sh-oil rosin soap at 5 g/L post-treatment irrigation is used to move the
also gave the highest protection from mealybugs insecticide in the root zone. Single application of
(Katke 2008). Application of spirotetramat 6 imidacloprid in spring through drip irrigation
.oz/acre + adjuvant (Vintre 0.25 %v/v) and 44 systems at rates of 0.75 g a.i. or higher per plant
.oz/acre (spray volume 137 gal/ac) were able to is known to reduce the mealybug abundance by
reduce the bunch infestation with mealybugs to 3 more than 99 % during the entire season and even
% and there was little to no honeydew in that for two seasons providing population pressures
treatment. Vintre dissolves mealybug-excreted remain low (Patricia Larrain 1999; Lo and Walker
wax to improve pesticide penetration, knock- 2011; Fu Castillo et al. 2004; Mansour et al.
down and control, deep within bark crevices. 2010; Mani et al. 2008). Imidacloprid provides
Diafenthiuron @ 8001600 g/ha recorded the 3060 % reduction in cluster damage when
340 M. Mani and U. Amala
applied through furrow irrigation (Daane et al. Table 38.2 List of insecticides recommended to control
mealybugs
2008a, b, c). In the furrow method, the vineyards
are prepared by ploughing a furrow area to Chemical Reference
expose the surface roots, followed by 1-day Buprofezin Muthukrishnan et al. (2005);
pretreatment irrigation. Imidacloprid is applied Mani et al. (2008); Bentley
et al. (2008); Daane et al.
into the furrows using the herbicide spray rig, and (2008a, b, c)
the application is followed by 1-day post-treat- Silwet L-77 Tipping et al. (2003)
ment irrigation. In the furrow method, there is a Imidacloprid Patricia Larrain (1999);
more widespread root zone that makes the deliv- Bentley et al. (2008);
ery of insecticide to the entire root zone difcult Gonzalez et al. (2001);
Daane et al. (2008a, b, c);
and results in a more dilute application and poor
Sunitha et al. (2009)
uptake of applied imidacloprid. Irrigation of both Azadirachtin Verghese (1997); Mani et al.
pre- and post-imidacloprid application is critical (2008)
and this is very difcult to properly manipulate Nimbecidine Koli (2003)
with furrow irrigation system. Neem-seed kernel Balikai (1999a); Koli (2003)
Soil application of granular insecticides, extract (NSKE)
namely phorate, carbofuran, thiodematon, fen- NSKE + soap powder Katke (2008)
sulfothion or bendiocarb (6 or 10 kg a.i./ha), once Neem oil Beevi et al. (1992)
after each pruning, was found to be ineffective in Neem oil + soap Katke (2008)
powder
reducing the mealybug infestation. However, one
Petroleum oil Michelakis and Hamid
application of granular insecticide aldicarb @ 50
(1995)
g/vine around the base of the plant at the time of
Fish-oil rosin soap Reddy and Narayana (1986)
October pruning protected the bunches com- Parathion Grimes and Cone (1985a);
pletely from mealybug infestation for 34 months AliNiazee and Stafford
(Anonymous 1984; Mani and Thontadarya 1991). (1972)
Though it is an excellent chemical for the mealy- Methyl parathion AliNiazee and Stafford
bug control in both April- and October-pruned (1972)
crops, the time of application (to be applied Permethrin Grimes and Cone (1985a)
immediately after pruning) is a critical factor, and Malathion Grimes and Cone (1985a);
Su and Wang (1988);
the farmers may not adhere strictly to the applica- Baskaran et al. (1999)
tion time and may apply the chemical to the vines Methidathion, AliNiazee and Stafford
particularly in the fruiting stage (many growers supracide (1972)
become aware of the damage at that stage, which Dimethoate AliNiazee and Stafford
will result in high residue problem). There are (1972); Shreedhar Rao et al.
(1988); Su and Wang (1988);
many restrictions in using aldicarb in the vine-
Baskaran et al. (1999); Sazo
yard ecosystem in India. A list of insecticides et al. (2008)
recommended to control mealybugs is given in Dimethoate + Fish-oil Katke (2008)
Table 38.2. rosin soap
Methomyl Mani et al. (2008); Balikai
(1999a); Raguraman and
Premalatha (2006)
38.5.4 Monitoring of Ants
Chlorpyriphos Mani et al. (2008); Bentley
et al. (2008); Hatta and Hara
Ants are known to attack the predators of scales (1992)
and mealybugs while attending to the pests. Dichlorvos Mani et al. (2008); Balikai
Therefore, it is necessary to check the activity (1999a); Shreedhar Rao
of ants prior to the release of Cryptolaemus. et al. (1988); FIP (1982)
General ant control measures like destruction of Dichlorvos + sh-oil Mani (1990); Beevi et al.
rosin soap (1992)
ant holes and ant nests, application of sticky
(continued)
38 Fruit Crops: Grapevine 341
A number of natural enemies are known to ists. Few fungal pathogens are also known to
attack vine mealybugs throughout the world. infect mealybugs in nature. However, mealybugs,
Many of the parasitoids are mealybug species being sessile insects, are more amenable to bio-
specic, while most of the predators are general- logical control.
342 M. Mani and U. Amala
of the year (off season) facilitating the non- dactylopii may not be useful in controlling the
interference with the predator. Almost 3 months pink mealybug since it is present already in
are available for the predator (JulySeptember) nature and has attained biotic balance. Anagyrus
to clear the residual population of mealybugs kamali (Moursi), an encyrtid parasitoid, caused
present on the vines. A mean of 97.67 % reduc- 8090 % reduction in population density of pink
tion in the bunch infestation was obtained with hibiscus mealybug at release sites in Egypt,
the release of C. montrouzieri in July (Fig. 38.3). Caribbean Islands and the United States. This
Depending upon the severity of infestation, the parasitoid is to be tried against M. hirsutus in
beetles have to be released. A release rate of vineyards in India.
5,000 beetles/ha is recommended to suppress the Lecanicillium (Verticillium) lecanii
pest population. Two to three releases are to be (Zimmerman) and Metarhizium anisopliae
made annually depending upon the severity of (Metch) are known to cause mortality of mealy-
pest infestation. The releases have to be made bugs (Katke 2008; Humber and Soper 1981). The
early in the season. The rst generation develops pathogen V. lecanii (Zimmerman) was isolated
from the released beetles. The second generation from whiteies and developed as a biopesticide
denitely brings down the pest population. As a named as Phule bugicide at Mahatma Phule
prerequisite for release, spraying of insecticides Krishi Vidyapeeth, Rahuri, Maharashtra, India,
has to be discontinued for 23 weeks prior to the for the control of mealybugs. A dosage of 20 g
release of the predator. It is better to release less formulated material/10 L of water is recom-
number of beetles at many places in unit area of mended to control the mealybugs. Two to three
the vineyard rather than more number of beetles sprays at 15-day intervals in rainy season are
at few places. needed. Addition of milk powder 5 g/10 L water
Anagyrus dactylopii is the naturally occurring helps to improve the control of mealybugs. Foliar
parasitoid on Maconellicoccus hirsutus in India. sprays of fungal pathogens, namely Beauveria
Inundative augmentation by ooding the chosen bassiana (Bals.) Vuill and Metarhizium aniso-
area with large numbers of particular natural pliae, in the rainy season under humid conditions
enemy is intended to exert rapid control of the were also found to infect the mealybugs.
pest in the present generation and prevent or Verticillium lecanii WP (Wettable Powder) @ 0.3
bring down the possible damage to host. % was found to be best against nymphs and
Conservation of the native A. dactylopii through adults of grape mealybug in Maharashtra (Koli
parasite-friendly insecticides like dichlorvos or 2003). The fungus was known to cause 80 %
buprofezin is to be done. Inundative release of A. mortality of some sucking pests at 2 105 cfu/mL
dose in Maharashtra state within two weeks Province and Georgia, P. citri was effectively con-
(Jayachakravarthy 2002). Fish-oil rosin soap (0.5 trolled by the release of C. montrouzieri (Aykac
%) with V. lecanii (0.4 %) was the safest and and Erguder 1972) (Dzhiviladze 1979).
most suitable treatment against grapevine mealy- The main parasite of P. citri infesting grapes is
bug, M. hirsutus (Shelke 2001). Anagyrus pseudococci (Girault), which occurs in
the south of European Russia and in Soviet
Central Asia and which destroys up to 75 % of
38.7 Planococcus citri (Risso) the coccid population in areas not treated with
insecticides (Niyazov 1969). Leptomastix dacty-
Besides Spain and Brazil, Planococcus citri lopii (Howard) is an effective encyrtid parasitoid
(Risso) is injurious to grapevine in the Soviet of citrus mealybug Planococcus citri (Risso).
Union. The main parasite of P. citri is Anagyrus The parasitoid can be multiplied on the
pseudococci (Girault), which occurs in the south laboratory-bred P. citri, P. lilacinus (Cockerell)
of European Russia and in Soviet Central Asia and P. minor (P. pacicus) (Cox). L. dactylopii
and which destroys up to 75 % of the mealybug was recovered in large numbers from P. citri
population in areas not treated with insecticides. infesting wine grapes in Maharashtra. It gives
Allotropa mecrida (Walker), the second most scope of utilizing L.dactylopii to control P.citri in
important parasite, was reared from the mealy- vineyards in Maharashtra and Karnataka. It has
bug in Turkmenia, and in Georgia it is responsi- given excellent control of P. citri in citrus in India
ble for up to 20 % parasitism. In 1960, (Krishnamoorthy and Singh 1987) and guava
Leptomastidea abnormis (Girault) and ecosystems in India (Mani 1994). Alternatively,
Leptomastix dactylopii (Howard) were intro- inundative releases of the local Coccidoxenoides
duced into Georgia and subsequently into perminutus can be done to suppress P. citri since
Turkmenia from the United States. In it can be multiplied easily in large numbers and it
Transcaucasia and Soviet Central Asia, Thysanus is a major parasitoid of P. citri in citrus ecosys-
(Chartocerus) subaeneus (Forster) is responsible tem in India (Mani 1994).
for up to 1820 % parasitism of Allotropa
mecrida. Others are Coccinella septempunctata
(Linnaeus), Hyperaspis polita (Weise), Scymnus 38.8 Pseudococcus longispinus
apetzi (Mulsant), S. subvileosus (Goeze), S.
bipunctatus (Kug.) and S. biguttatus (Mulsant), Currently, Ps. longispinus infests a small number
which were noted in Turkmenia. The larvae of of vineyards in Californias coastal region.
Leucopis (Leucopomya) alticeps (Czerny) and Recent surveys found Trachelomonas sydneyen-
Chrysoperla carnea (Stephen) destroy virtually sis (Playfair), Tetracnemus peregrina (Compere),
all stages of the mealybug (Niyazov 1969). Acerophagus angelicus (Howard), Anagyrus
Cryptolaemus montrouzieri has given excellent pseudococci, Leptomastidea abnormis (Girault),
control of P. citri in several ecosystems in many Leptomastix dactylopii (Howard) and
countries. Extensive eld trials were conducted in Coccidoxenoides perminutus (Girault) attacking
Karnataka and Maharashtra in India on the use of this mealybug (Daane et al. 2008a).
Cryptolaemus for the control of P. citri mealybug
on grapevine. Cryptolaemus montrouzieri proved
to be very useful in suppressing the mealybug in 38.9 Planococcus cus
the grape gardens (Mani and Krishnamoorthy
2008). In USSR, C. montrouzieri was one of the As many as 20 natural enemies were recorded on
most effective predators introduced into Black Sea P. cus infesting grapes. Natural enemy complex
Coastal area for the control of P. citri in the vine- consists of the parasitoids, namely Angyrus sp.,
yards (Niyazov 1969). In the vineyards of Tokat Coccidoxenoides perminutus (Girault),
38 Fruit Crops: Grapevine 345
in the rainy season (JulyAugust) can also be Balikai RA (1999a) New record of alternate host plants of
grape mealy bug. Insect Environ 5(2):81
given.
Balikai RA (1999b) Seasonal incidence of grapevine,
One or two applications of dichlorvos 76 % mealybug in North Karnataka. Insect Environ
EC (2 mL/L) from mid-February to the rst 4(4):148149
week of March, if necessary, depending upon Baskaran RKM, Lakshmi LG, Uthamasamy S (1999)
Coccids and their management in guava intercropped
the incidence of mealybugs and time of har-
with coconut. Pest Manage Hortic Ecosyst
vesting or one jet spray of water can also be 5(1):2831
given on the bunches if the mealybugs are still Batra RC, Brar SS, Khangura JS, Dhillon WS (1987) A
present just prior to harvest to dislodge the new record of Planococcus pacicus Cox.
(Pseudococcidae: Hemiptera) as a pest of grapevine in
mealybugs.
India. Punjab Hortic J 27(34):250251
Beevi ND, Janarthanan R, Natarajan K (1992) Efcacy of
some insecticides against Maconellicoccus hirsutus
(Green) on mulberry. J Insect Sci 5(1):114
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Su TH, Wang CM (1988) Control measure for the citrus on table grapes on the mealybug parasitoid
mealybugs and latania scale insects of grapevine. Coccidoxenoides peregrinus (Timberlake)
Plant Prot Bull (Taiwan) 30(3):279288 (Hymenoptera: Encyrtidae). S Afr J Enol Viticult
Subba Rao BR, Sangwar HS, Abbasi OA, Singh Y, Ksheer 20:3134
Sagar AM (1965) New records of hymenopterous Walton VM, Pringle KL (2001) Effects of pesticides and
parasites of Nipaecoccus vaster Maskell (Homp. fungicides used on grapevines on the mealybug preda-
Coccidae) a serious pest of citrus from India. Indian tory beetle Nephus boschianus (Coccinellidae,
J Entamol 22:109110 Scymnini). S Afr J Enol Viticul 22:107110
Sunitha ND, Jagginavar SB, Biradar AP (2009) Walton VM, Pringle KL (2004) A survey of mealybugs
Bioefcacy botanicals and newer insecticides against and associated natural enemies in vineyards in the
grapevine mealybug, Maconellicoccus hirsutus Western Cape Province, South Africa. S Afr J Enol
(Green). Karnataka J Agric Sci 22(3):710711 Viticul 25:2325
Swart PL, Barnes BN (1975) Improved mealybug control Walton VM, Daane KM, Pringle KL (2004) Monitoring
on table grapes with a quick-breakdown insecticide. Planococcus cus in South African vineyards with sex
Decid Fruit Grow 25(1):4, 68 pheromone-baited traps. Crop Prot 23:10891096
Swart PL, Barnes BN (1976) Mealybug on table grapes: a Whitehead VB (1961) Insect enemies of the mealybug on
most effective insecticide for the control of the vine grapevines. Decid Fruit Grow 10:123127
mealybug, and the economic implications of three- Williams DJ (2004) Mealybugs of southern Asia. The
and four-spray programs. Decid Fruit Grow Natural History Museum/Southdene SDN. BHD,
26(10):378382 London/Kuala Lumpur, 896 p
Tejkumar S, Aftab Ahmed M, Dhramaraju E (1977) Willink MCG, Scatoni IB, ETerra AL, Frioni MI (1997)
Occurrence of the mealy bug, Pseudococcus spp. A Cochinillas harinosas (Homoptera, Pseudococcidae)
serious pest of grapevine around Hyderabad. Indian que afectan plantas cultivadas y silvestres en Uruguay.
J Entamol 39:189190 Agrociencia Montevido 1:96100
Thiem H (1925) The most important coccids injurious to Wunn H (1928) Observations on coccids Bad. 81. Z
orchards and vineyards in Germany Biol. Reiehsanst, Angew Ent ii(6):321322
Land-u-Forstw. Flugbl 77:4 Zhang AJ, Amalin D, Shirali S, Serrano MS, Franqui RA,
Tipping C, Bikoba V, Chander GJ, Mitcham EJ (2003) Oliver JE, Klun JA, Aldrich JR, Meyerdirk DE,
Efcacy of Silwet L-77 against several arthropod pests Lapointe SL (2004) Sex pheromone of the pink hibis-
of table grape. J Econ Entomol 96(1):246250 cus mealybug, Maconellicoccus hirsutus, contains an
Tranfaglia A, Viggiani G (1981) Problems of integrated unusual cyclobutanoid monoterpene. Proc Natl Acad
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Transfaglia A (1976) Considerationson the morphological bug, Phenacoccus hystrix (Bar) in the vine region of
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(Homoptera : CoecoideaL In Alii XI Congresso XVII:67102
Fruit Crops: Citrus
39
C.N. Rao, V.J. Shivankar, K.J. David, M. Mani,
and A. Krishnamoorthy
Table 39.1 List of mealybug species recorded on citrus in different regions of the world
Species Region/Country References
Crisicoccus matsumotoi (Siraiwa) India Williams (2004)
Philippines Williams (2004)
Dysmicoccus brevipes (Cockerell) India Williams and Watson (1988)
Australia Ben-Dov (1994)
Middle East Ben-Dov (1994)
Columbia Kondo et al. (2008)
USA Ben-Dov (1994)
Japan Borchsenius (1956); Ben-Dov (1994)
Russia Borchsenius (1949); Borchsenius (1956)
Argentina Granara de Willink (1991)
Crisicoccus chiponensis (Takahashi) Taiwan Williams (2004)
Crisicoccus matsumotoi India Williams (2004)
(Siraiwa)
Dysmicoccus lepelleyi (Betrem) Ben-Dov (1994)
Dysmicoccus debregeasiae (Green) India, Bangladesh, Williams (2004)
Nepal, Sri Lanka
Dysmicoccus neobrevipes Beardsley India Williams and Watson (1988),
Williams and Granara de Willink (1992),
Williams (2004)
Australia Ben-Dov (1994), Williams and Watson
(1988)
Brazil Williams and Granara de Willink (1992),
Ben-Dov (1994)
Pakistan Williams (2004)
Philippines, Sicily Ben-Dov (1994)
Ferrisia consobrina Williams and Australia, Ethiopia, Ben-Dov (1994)
Watson Neotropical and Pacic region
Ferrisia terani Argentina Ben-Dov (1994)
Williams and Granara de Willink
Ferrisia malvastra (McDaniel) India Williams and Watson (1988), Williams
and Granara de Willink (1992), Williams
(2004)
Australia Williams and Watson (1988)
South Africa Ben-Dov (1991), Ben-Dov (1994)
Argentina Williams and Granara de Willink (1992);
Ben-Dov (1994)
Ferrisia virgata (Cockerell) India Ali (1968), Williams and Watson (1988),
Williams and Granara de Willink (1992),
Williams (2004), Mani and
Krishnamoorthy (2008)
Australia Williams (1985), Ben-Dov (1994)
South Africa Ben-Dov (1994)
Columbia Kondo et al. (2008)
Bangladesh Williams (2004)
USA Ben-Dov (1994)
Argentina Granara de Willink (1979/1991)
(continued)
39 Fruit Crops: Citrus 355
39.2 Damage with mealybugs and sooty mould lose the mar-
ket value (Bindra 1970; Rao et al. 2001). In
Nymphs and adult mealybugs suck the sap from recent years, the incidence of mealybugs caus-
shoot, leaf, bark, stem, owers and fruits. Fruit ing losses to citrus has been reported from pen-
production is often adversely affected due to insular India (Krishnamoorthy and Singh
the mealybug damage in citrus. There is an 1987), the north-east region (Pathak et al. 1999;
upward trend in the build-up of mealybugs Shylesha and Pathak 1999) and Punjab (Arora
adversely affecting the growth of citrus plants et al. 1999) on different citrus cultivars.
of all ages from young seedlings to grown-up Maconellicoccus hirsutus is known to inject
trees, which occasionally attain epidemic forms toxic saliva into the plant while feeding, which
in citrus. In addition, the sticky honeydew results in malformation of leaf and shoot
excreted by the mealybugs serves as a substrate growth, stunting and occasional death. Leaves
for the growth of sooty mould interfering with show a characteristic curling, while heavily
photosynthesis (Butani and Srivastava 1999). infested plants have shortened internodes lead-
The mealybug infestation on citrus plants ing to resetting or a bunchy top appearance
ranged from 38 to 65 % in India. Fruits covered (Mani and Krishnamoorthy 1999).
Howard and Anagyrus pseudococci (Girault) are Nephus sp., Chrysopa sp., Micraspis cardoni
common wasps of P. citri. Some mealybugs are (Wse.), Pseudaspidimerus uttami Kap.,
susceptible to infections by the entomopatho- Cryptolaemus montrouzieri Muls. and Spalgis
genic fungus, Entomophthora fumosa. Common epeus Westwood were recorded on mealybugs
predators include the brown lacewing, in Kodagu, Karnataka (Singh 1990).
Sympherobius barberi (Banks), and the green Coccidoxenoides perminutus (Timberlake),
lacewing, Chrysopa lateralis Gurin, trash bugs, Mallada boninensis (Okamoto), Plesiachrysa
syrphid y larvae, and the scale-eating caterpil- lacciperda (Kimmins), Anisochrysa basalis
lar, Laetitia coccidivora, the Australian ladybird Walker and Chrysoperla carnea (Steph.)
beetle, Cryptolaemus montrouzieri Mulsant, the (Krishnamoorthy and Mani 1988a,1989a, b;
little mealybug-eating lady beetle, Decadiomus Krishnamoorthy and Mani 1988b) were
bahamicus (Casey) and the pictured ladybeetle, recorded in Karnataka. In Assam, C. montrouz-
Scymnus avifrons Melsheimer, which feed pri- ieri and Entomophthora fumosa Speare were
marily on mealybugs. Two other lady beetles, observed on P. citri (Chowdhury and Majid
Chilocorus stigma (Say) and Olla abdominalis 1954). In central India, Cacoxenus perspicax
var. plagiata (Say), occasionally feed on mealy- and Promuscidea unfasciativentris were
bugs (Muma 1954). observed on P. citri. C. montrouzieri is known to
In India, Anagyrus sp., Blepyrus insularis consume about 3,300 eggs of P. citri. The eggs
(Cam.), Diversinervus sp., Tetrastichus sp., as well as other stages of mealybug are essential
Microterys sp., Cryptochaetum sp., Scymnus in the diet for successful development of the
coccivora Ayyar, Pullus pallidicollis Mst., predator (Oncuer and Bayhan 1982).
sprays should be applied immediately after most Chemical control of Pseudococcus longispi-
of the eggs have hatched to prevent the crawlers nus was most effective if sprays were applied
from hiding themselves in crevices when the mealybugs were in the dispersive
The intervention threshold for P. citri on crawler stage and when the food-plant afforded
Nagpur mandarin was reported to be 510 % the least shelter. A two-spray programme with
infested fruits in summer and 15 % infested fruits applications in August and late November effec-
in autumn (Shivankar and Shyam Singh 2000). tively controlled a dense infestation on citrus. An
Spraying of 1.5 ml dimethoate + 2.5 ml kerosene overall pest management programme has been
oil in 1 l water or 1 ml carbaryl + 1 ml kerosene developed for citrus, in which all the insect pests
oil or 2 ml malathion in 1 l water checks mealy are controlled by combinations of natural ene-
bugs effectively. Spraying with chlorpyriphos mies and insecticides as required. Outbreaks of
0.05 %, carbaryl 0.1 % or fenitrothion 0.05 % Ps. longispinus and other secondary pests are
(Jadhav et al. 1997; Jadhav and Pujari 1999) and controlled by sprays of aminocarb and metho-
also with 2 ml dichlorvos + 25 g sh oil rosin myl. These two insecticides prevented the resur-
soap in 1 l water resulted in 75 % of reduction in gence of the mealybug in the subsequent
mealybug population. Since the pest harbours generation that occurred when malathion was
under the loose bark, debarking by treating with used (Furness 1977). N. viridis is susceptible to
chlorpyriphos and methyl parathion (both at 4 treatments of methidathion and chlorpyriphos.
ml/l) helps in minimizing the pest population On heavy infestations, repeated treatments are
(Mani and Krishnamoorthy 1996). The settling of essential (Bar Zaki et al. 1988). Effective control
crawlers on the plant was reduced by swabbing it was achieved by Mospilan (acetamiprid 20 solu-
with carbaryl (1 %), used for trunk borer man- ble powder (SP)) 0.05 % and 0.075 % sprayed on
agement (Shylesha and Pathak 1999). small fruits or by 0.3 % chlorpyrifos at both
Chlorpyriphos performs very well against the cit- stages of the development of N. viridis (Gross
rus mealybug; but, since this product is especially et al. 2000).
harmful to parasitoids, it is not considered to
have a good t in the integrated pest management
(IPM) programmes, where parasitoid conserva- 39.6.4 Insect Growth Regulators
tion is emphasized.
Citrus oil mixed with chlorobenzilate is effec- Buprofezin, an insect growth regulator, showed
tive against rst-instar nymphs of the mealybugs strong ovicidal activity resulting in more than 80
in an integrated control programme (Meyerdirk % inhibition of egg hatch and 9199 % nymphal
et al. 1981). On grapefruit in the Spanish Province mortality of P. citri (Mendel et al. 1991). It was
of Castellon against Planococcus citri, spot treat- an effective treatment and should be considered
ments on the trunks and main branches of the trees for the citrus mealybug control to avoid destruc-
provided complete control. The products tested tion of parasitoids.
contained 50 % fenitrothion, a mixture of 50 % The addition of narrow-range crop oil, NR-415
fenitrothion + 40 % dimethoate, 50 % omethoate, at 1.0 gal/ac, appeared to be benecial for initial
40 % methidathion, 40 % dimethoate or 50 % mealybug knock-down, especially for the slower-
dichlorvos (Limon de la Oliva et al. 1972). In acting insecticides such as buprofezin. ZR-777
California, dimethoate remained effective for 3 (prop-2-ynyl 3,7,11-trimethyl-(2E,4E)-dodecadi-
months against Planococcus citri (Risso), a seri- enoate), which exhibits high morphogenetic
ous pest attacking the greenhouse crops. activity against the species of Homoptera, gave
Soil injection of imidacloprid (Admire) at 16 good control of nymphs in all instars receiving
and 32 oz/ac appeared to have a very good activ- 0.01 % sprays (Staal et al. 1973). In grapefruit
ity against P. citri (Kerns et al. 2004). orchards, 0.0250.10 % kinoprene, 0.025 %
39 Fruit Crops: Citrus 363
CGA-13353 [ethyl 3-methyl-4-[4-(phenylmethyl) nymphs and least effective against adults (French
phenoxy]-2-butenoate] and 0.0250.10 % and Reeve 1979). In Egypt, the insect growth
epofenonane (Ro 103108) gave good control of regulator methoprene (Altosid) applied by spray-
citrus mealybugs compared to the control that ing on the infested citrus leaves at 0.01 and
was obtained with 0.06 % dimethoate. In tests on 0.05 % gave satisfactory suppression of popula-
the efcacy against specic life stages, both tions of the citrus pest Planococcus citri.
insect growth regulators and conventional insec- Concentrations of 0.1 and 0.5 % gave 100 % sup-
ticides were most effective against rst-instar pression of mealybugs (Hamdy 1984).
Leptomastix dactylopii
California Florida
Biological control efforts against the citrus C. montrouzieri was introduced into Florida in
mealybugs, chiey P. citri, in California started 1930 for the control of P. citri on citrus and bulbs.
with the introduction of C. montrouzieri in 1891 Despite permanent establishment, it failed to sur-
1892. It readily cleaned up the infestations but vive in sufcient numbers from year to year for
subsequently proved to be generally slow in adequate control (Bishop 1931; Watson 1932;
response and unable to survive in winter eventu- Muma 1954).
ally persisting only along the coast (Clausen
1915). Complete control of P. citri in some Hawaii
orchards in California was obtained with the con- P. citri was partially controlled by C. montrouz-
tinued liberations of large numbers of C. mon- ieri that was accidentally introduced in Hawaii
trouzieri (Smith 1919). After being colonized in (Bartlett 1977).
citrus mealybug-infested orchards in California,
these ladybird beetles sometimes showed remark- USSR
able ability to destroy many kinds of mealybugs C. montrouzieri was used to control P. citri in
(Hoyt 1912). Later, great interest was stimulated Black Sea coast (Rubtsov 1954). In the Soviet
in this method of periodic colonization. Union, Planococcus citri (Risso) is injurious to
Subsequently, many insectaries were established over 20 species of plants, including Citrus. The
and periodic colonization of Cryptolaemus main parasite of P. citri is Anagyrus pseudococci
reached a peak in the 1920s against citrus mealy- (Gir.), which occurs in the south of European
bugs. Ten beetles per tree during summer were Russia and in Soviet Central Asia and which
adequate for the control of most of the infesta- destroys up to 75 % of the coccid population in
tions of the mealy bugs. More than 40 million areas not treated with insecticides. Allotropa
beetles were released over some 50,000 acres of mecrida (Wlk.), the second most important para-
citrus during 19261927 (Essig 1931). In 1928, site, was reared from the coccid in Turkmenia in
over 42 million beetles were released in citrus in 1967 and is responsible for up to 20 % parasitism
California. An average of 23 adults were released in Georgia. In 1960, Leptomastidea abnormis
per tree and over one million trees received lib- (Gir.) and Leptomastix dactylopii. How were
erations (Beckley 1956). In addition, the number introduced into Georgia and subsequently into
of outbreaks of P. citri was reduced due to the Turkmenia from the United States (Niyazov
presence of Cryptolaemus (Clausen 1956). The 1969).
outstanding reduction in citrus mealybug was
related to the peak period of Cryptolaemus activ- France
ity. The predator was more active in late April C. montrouzieri was introduced in France from
and the activity started decreasing in June as the California in July 1918 against P. citri (Turinetti
mealybug population declined (Bartlett 1957). It 1921). The predator became established and pro-
was, however, rated as partial control (Debach duced a marked reduction in the numbers of P.
and Hagen 1964). In the later years, the citrus citri. But its overwinter survival was low
mealybug was kept under check by the periodic (Marchal 1921; Anonymous 1922; Poutiers
colonization of C. montrouzieri along with the 1922; Marchal and Pussard 1938). An outbreak
encyrtid Leptomastix dactylopii Howard of P. citri at Cap dAntibes was checked by C.
(Beckley 1960). The Brazilian encyrtid parasit- montrouzieri without any liberation of the preda-
oid, Leptomastix dactylopii How., was utilized in tor at that time. It might be due to the develop-
the suppression of the mealy bug Planococcus ment of adapted strain from the beetles released
citri in USA (Fisher 1963). earlier in the 1920s (Pussard 1938).
39 Fruit Crops: Citrus 365
since it did not establish permanently (Duran declined from 313.84/plant in August 2005 to
1944; Gonzalez and Rojas 1966). 2.63/plant in October 2005 following the release
of C. montrouzieri at 30 larvae/plant in August
Australia 2005 (Mani and Krishnamoorthy 2008). Several
The predator was brought from New South Wales green lace wings preying on mealybugs have
in 1902 to Eastern Australia for the control of the been reported from India (Krishnamoorthy and
citrus mealybugs. It was readily established and Mani 1988a).
effective control was obtained. Its introduction is The encyrtid parasitoid Coccidoxenoides
regarded as the outstanding biological control peminutus played a dominant role in the suppres-
success (Wilson 1960). sion of P. citri on acid lime and lemon in India
(Mani 1994). The parasitoid was multiplied on
South Africa 510-day-old laboratory-bred P. citri. Several
The liberations of C. montrouzieri were made in plant products and deltamethrin were found to be
the citrus mealybug-infested orchards from early safer to C. perminutus. The exotic parasitoid
January, and good control was obtained in every Leptomastix dactylopii was imported from West
instance (Bishop 1931). The biological control of Indies to India during 1983 for trials against P.
the citrus mealybug by C. montrouzieri was rated citri (Nagarkatti et al. 1992). Releases of L. dac-
as complete control in South Africa (Greathead tylopii gave excellent control of P. citri causing
1971). Allotropa kamburovi sp. n., recovered up to 100 % parasitism in citrus orchards in
from Planococcus citri (Risso), is derived from Karnataka (Manjunath 1985; Krishnamoorthy
citrus in Western Transvaal, South Africa and Singh 1987; Nagarkatti et al. 1992;
(Annecke and Prinsloo 1977). Cryptolaemus Krishnamoorthy 1990). L. dactylopii released in
montrouzieri is used in augmentation programme April 1984 was recovered in large numbers from
in the control of P. citri on citrus in South Africa P. citri infesting acid lime and lemon in 1991
(Moore and Hattingh 2004). 1992 (Mani 1994). Later L. dactylopii was recov-
ered from P. citri infesting several horticultural
Palestine crops (Krishnamoorthy and Mani 1989c).
No practical success in the control of P. citri was Dichlorvos, dicofol, several fungicides and plant
observed with C. montrouzieri (Bodenheimer products are safer to L. dactylopii (Mani et al.
1928, 1951). 1993). When L. dactylopii and C. perminutus
were found together, with the latter one playing a
India dominant role in suppressing P. citri. However,
During 19631965, the release of the coccinell- L. dactylopii is known to prefer the late-nymphal
lid, C. montrouzieri, did not result in establish- instars of P. citri (Krishnamoorthy 1988), while
ment in the citrus orchards, located 20 miles C. perminutus is known to attack preferentially
away from Gauhati, India. It might be due to the the early-nymphal instars (Krishnamoorthy and
activity of the ant Oecophylla smaragidina Mani 1988b). L. dactylopii had not displaced the
(Narayanan 1957). A release rate of ten beetles local C. peminutus in Bangalore. Under this situ-
per Coorg mandarin orange tree was suggested ation, C. perminutus was largely responsible for
for the control of citrus mealybug in Karnataka the control of P. citri in both the orchards. Similar
(Singh 1978). Following the release of C. mon- control of P. citri on grapefruit was achieved in
trouzieri at 2000/acre (ac) on acid lime plants, the April when C. perminutus was abundant in
population of mealybug (P. citri) was declined October in Texas (Dean et al. 1971). The same
from 126.64 in August 2003 to 0.4/plant in parasitoid has also been used to control P. citri in
November 2003. A mean of 99.68 % reduction in Peru and Bermuda (Bartlett 1977). In India, the
the mealybug population on acid lime was mealybug infestation was rst noticed in the sec-
achieved by the predator within 3 months of its ond week of February on lemon in India. The
release (Mani and Krishnamoorthy 2007). In the mean number of mealybugs per ve shoots was
pummelo orchard, the population of P. citri 1342.4, and initial samples, collected on 16
39 Fruit Crops: Citrus 367
February 1991, yielded both L. dactylopii and C. main pest of citrus, was controlled by the natural
perminutus. Both the parasitoids were active up enemies including C. montrouzieri (Ozkan et al.
to the second week of May. C. perminutus was 2001).
always found to emerge in larger numbers than L.
dactylopii from all the samples collected from Morocco
February to May. It was particularly abundant in In Morocco, P. citri is a major pest of citrus
March/April, and a mean maximum of 318.5 orchards. The predator C. montrouzieri was used
parasitoids emerged from the samples collected to control the mealybug pest (Abdelkhalek et al.
on 1 April 1991. In the case of L. dactylopii, a 1998).
mean maximum of 49.2 adults were recovered
from the samples collected on 2 March 1991. Bermuda
There was a marked reduction in the mealybug The encyrtid parasitoid L. dactylopii has also been
population, which was negligible in the second used to control P. citri in Bermuda (Bartlett 1977).
week of May.
39.6.5.2 Japanese Mealybug
Cyprus Planoccoccus kraunhiae
C. montrouzieri was imported from California An isolated infestation of P. kraunhiae on citrus
and releases were carried out in 1954 and 1960 disappeared following the release of C. mon-
resulting in temporary establishment, but the bee- trouzieri in Southern California (Smith and
tles did not survive the winter (Greathead 1976). Armitage 1931).
The Brazilian encyrtid parasitoid, Leptomastix
dactylopii How., has been utilized in the suppres- 39.6.5.3 Oriental Mealybug
sion of the mealy bug Pl. citri (Krambias and Planococcus lilacinus
Kontzonis 1980). The oriental mealybug appeared on 3-year-old
acid lime plants at Indian Institute of Horticultural
Greece Research (IIHR) farm, Bangalore in September
P. citri disappeared in 1933 as a pest after the 1998. The initial sampling done on 19th
introduction of the predator from Spain (DeBach September did not yield any natural enemy. Since
and Argyriou 1967; Pelakasis 1974). Recoveries C. montrouzieri is known to feed on P. lilacinus,
were also made following the introductions in releases of C. montrouzieri were made on 20th
1965 and 1969 (Argyriou 1974). September and 10th October at 20/plant. Prior to
the release of the predator, ants attending the
Brazil mealybugs were checked. A mean of 160.50
C. montrouzieri was used as a biological control mealybugs/shoot was observed when the study
agent against P. citri in Brazilian citriculture was initiated. The mealybug population had
(Gravena 2003). The Brazilian encyrtid parasit- started declining following the release of C. mon-
oid, Leptomastix dactylopii, has been utilized in trouzieri. It was found in negligible numbers by
the suppression of the mealy bug Planococcus mid-November 1998 and ceased to be a problem
citri in several countries. from January 1999. The cecidomyiid T. coccidi-
vora was also observed but in negligible numbers
Turkey in India.
C. montrouzieri was introduced into Turkey and The mealybug (P. lilacinus) population was
compared with native races in the control of P. rst observed in April 1991. More than 35 % of
citri in citrus orchards. There were no signicant fruits were found infested with P. lilacinus. The
variations in the cold hardiness, prey consump- population of L. dactylopii ranged from 2.46 to
tion capacity and other biological characteristics 6.00, but T. indica was observed in large numbers
of native and introduced races of C. montrouzieri ranging from 14.45 to 96.00. Due to the action of
(Yigit and Canhilal 1998). In Turkey, P. citri, the both the parasitoids, especially T. indica, the
368 C.N. Rao et al.
mealybug population was reduced by the end of 1931). The application of burlap bands around
July. The mealybug did not appear up to the trunks of infested trees attracted the mealy-
December 1991 (Mani 1995b). The 5-day-old P. bugs to congregate beneath the bands to oviposit.
lilacinus was found suitable for the breeding of T. The bands also attracted the coccinellids, thereby
indica (Mani and Krishnamoorthy 1995). increasing the intensity of predation. In San
Francisco, serious infestations of P. fragilis were
39.6.5.4 Citriculus Mealybug invariably controlled by C. montrouzieri (Smith
Pseudoccoccus cryptus 1928). In Abkhazia of USSR, complete control
(Ps. citriculus) of P. fragilis was secured in 1933 by releasing
In Israel, the imported C. montrouzieri released 1520 adults on severely infested tree or ten
in 1924 did not successfully establish adults on slightly infested tree. The coccinellid
(Bodenhiemer 1928). It was again released in thrived throughout the summer and autumn with-
1941 (Mason 1941) and 1958 (Rosen 1967), but out being affected by high humidity and torren-
there were no reports of permanent establish- tial rain or the maximum temperature of 35 C
ment. Four parasitoid species, Allotropa burrelli, (Stephanov 1935). Also in Chile, C. montrouzieri
A. convexifrons, Pseudaphycus malinus (from was utilized against P. fragilis. Pseudococcus
central Asia) and Anagyrus sawadai (from fragilis is recorded for the rst time in Italy on
Japan), were introduced into Israel during 1996 Citrus in the Province of Salerno in 1969. The
1997 against P. cryptus. Only Allotropa convexi- heaviest infestation was on the fruits of orange,
frons and Anagyrus sawadai successfully lemon and mandarin. P. fragilis appeared to be
parasitized P. cryptus and therefore were released controlled effectively by natural enemies, espe-
in the eld. So far, only A. sawadai has been cially the encyrtid Hungariella pretiosa (Timb.);
recovered. A considerable reduction in popula- Dendrocerus laevis (Ratz.) and an aphelinid of
tion densities of the pest has been recorded since the genus Coccophagus were also recovered
May 1998 in the major release site (Blumberg from the mealybug (Viggiani 1970).
et al. 1999a, 1999b).
Pseudococcus calceolariae
39.6.5.5 Citrophilus Mealybug Cryptolaemus montrouzieri, initially introduced
as a classical biological control agent for
Pseudococcus fragilis Pseudococcus calceolariae, in southern
(= P. citrophilus; P. gahani) California, was unable to survive in sufcient
Release of C. montrouzieri was made in 1916 in numbers to provide control without augmenta-
Southern California and it provided some control tion. C. montrouzieri is still commercially avail-
of P. fragilis. At Los Angles County, 4,000,000 able and being used in citrus to suppress the
adults were released over an area of 4,775 acres mealybug pests (Luck and Forster 2003).
at ten per tree. Although prolonged cool damp
weather at rst delayed the activities, the control 39.6.5.6 Obscure Mealybug
for the season later became entirely satisfactory. Pseudoccoccus obscures (Ps.
Over 350,000 adults were released in 3-weeks viburni)
time against P. fragilis in California. Infestation The mealybug responded well to periodic releases
appears to be less severe than the previous years of C. montrouzieri in citrus orchards of California
(Anonymous 1929). The rst release was sug- (Bartlett 1977).
gested between 1st and 15th April when the eld
temperature was 70 F and rainfall was low. 39.6.5.7 Grape Mealybug
Liberations were continued up to September
(Armitage 1929). The beetles released at rst Pseudococcus maritimus
increased to controlling numbers in the progeny C. montrouzieri did not give sufcient results in
of rst generation of adults. Banding citrus trees the control of P. maritimus on citrus (Timofeeva
beneted C. montrouzieri (Smith and Armitage 1979).
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Fruit Crops: Guava
40
M. Mani
40.1 Species foliage and fruit also become covered with large
quantities of sticky honeydew which serves as a
Mealybugs are reported to be injurious to guava medium for the growth of black sooty moulds.
in India, Bangladesh, Taiwan, South Africa, The sooty moulds and waxy deposits result in the
Egypt, Pakistan, Sri Lanka, Hawaii etc (Table reduction of photosynthetic area. Ornamental
40.1). plants and produce lose their market value.
Nymphs and adults suck the sap from leaves, stem In Taichung, Taiwan, populations of both nymphs
and fruits. In addition, the sticky honeydew and adults of Planococcus citri on guava were
excreted by the mealybugs serves as a substrate large in the cool dry months from November to
for the growth of sooty mould interfering photo- April and small in the warm wet months from
synthesis. Severe mealybug infestation causes July to September. Nymphal populations had 4 or
heavy economic losses (Mani and Krishnamoorthy 5 marked peaks between November and April,
1993). In the case of Maconellicoccus hirsutus, considered ideal times for insecticide applica-
the mealybug injects toxic saliva into the plant tions. There was a negative relationship between
while feeding which results in malformation of mealybug populations and temperature. Incessant
leaf and shoot growth, stunting, and occasional rainfall and heavy pruning of trees also had
death. Leaves show a characteristic curling, while adverse effects on populations (Liu and Chang
heavily infested plants have shortened internodes 1984). In India, the mealybugs are found through-
leading to rosetting or a bunchy top appearance out the year on guava plants but the population
(Mani and Krishnamoorthy 2001). Infestations of was found in greater numbers in summer months
Ferrisia virgata remain clustered around the ter- (February-June).
minal shoots, leaves and fruit, sucking the sap
which results in yellowing, withering and drying
of plants and shedding of leaves and fruit. The 40.4 Management
Application of insecticides also eliminates the the parasitoids and predators are able to suppress
important naturally occurring bioagents resulting the mealybugs on guava (Mani and
in the outbreak of mealybugs. On the other hand, Krishnamoorthy 1990a, 2007; Manjunath 1986).
Releases of the exotic parasitoid Leptomastix May to 1.42 in September, 1990. The mealybug
dactylopii were found to be highly effective population was negligible after September, 1990
against P. citri in guava orchards (Mani 1994). In to December, 1991 (Mani 1994).
a guava orchard in Bangalore North, initial sam- In another guava orchard also in Bangalore
ples collected on June 14, 1990 revealed the pres- North, a mean mealybug population of 1954
ence of exotic parasitoid L. dactylopii but at a was observed in February 1991. A total of
very low level. At the same time, the mean mealy- 2,000 L dactylopii was released in February,
bug population was 1,084. This was due to indis- 1991. The parasitoid was recovered only after
criminate application of insecticides like methyl the releases made in March. The samples col-
parathion, monocrotophos and BHC against P. lected between 23rd March and 10th May, 1991
citri. However, after the suspension of insecti- yielded a large number of adult L dactylopii
cidal sprays, the population of L. dactylopii (Table). However, the local natural enemies like
started increasing. It was found in large numbers C. Natural enemy complex of P. citri consisted
between 23rd July and 4th August, 1990. Natural of the encyrtid parasitoid, Coccidoxenoides per-
enemy complex of P. citri consisted of the encyr- minutus (Timberlake), the gregarious platygas-
tid parasitoid, Coccidoxenoides perminutus terid Allotropa sp. besides L. dactylopii. In
(Timberlake) and the gregarious platygasterid general, C. perminutus was observed in small
Allotropa sp. besides L. dactylopii. C. perminutus numbers in the present study. But, Allotropa sp.
was observed in small numbers, but, Allotropa emerged in large numbers in the late samples
sp. emerged in large numbers in the late samples collected in August-September, Chrysopa sp.
collected in August-September. Indigenous pred- and C. montrouzieri Muls. remained at a very
ators like the lycaenid Spalgis epeus Westwood low level throughout the study. P. citri once
and Chrysopa sp. were noticed in very low num- found in very high numbers in February, 1991
bers. Due to build-up of the population of the almost completely disappeared by the end of
parasitoids especially L. dactylopii, the popula- May 1991, and the mealybug was kept under
tion of P. citri gradually declined from 1,084 in check up to December, 1991 (Mani 1994).
Neem oil and pongamia oil (both at 4 %) are 10 days after the rst) (Hussain and Puttaswamy
recommended for the control of P. citri on guava; 1996).
they caused 93.23 and 89.39 % mortality of the At Taichung, Taiwan, insecticide treatments
pest, resp., 10 days after the second spray (applied were generally more effective against nymphs
382 M. Mani
than adults. Spray applications made two or three guava orchards (Krishnamoorthy and Mani
times at intervals of 710 days prior to popula- 1988). A single larva of M. boninensis was able
tion build-up gave the best control. Application to prey about 345, 490 and 560 nymphs of F. vir-
of omethoate, methidathion, formothion and gata, P. lilacinus and P. citri respectively (Mani
dimethoate at various rates gave effective control and Krishnamoorthy 1990b). Blepyrus insularis
of P. citri. However, treatment with mixtures of bred very well on 510 old nymphs of F. virgata
malathion with methidathion or formothion, each (Mani and Krishnamoorthy 1991). The key para-
applied at half the rate when used alone, gave sitoid A. advena could be conserved by the appli-
effective and economic control, without being cations of diazinon, phosalone and dichlorvos
phytotoxic (Liu and Chang 1984). (Mani and Krishnamoorthy 1992).
Introduction of parasitoids gave improved Cryptolaemus montrouzieri was released
biological control of P. citri in a large glasshouse against F. virgata on guavas in the guava orchards
stocked with guava plants in the UK, supple- in India in 1987. C. montrouzieri along with the
menting that achieved by the coccinellid predator local natural enemies Aenasius advena Compere
Cryptolaemus montrouzieri. Following parasit- and Scymnus coccivora Ayyar effectively con-
oid release, there was evidence of pest population trolled the mealybugs within 50 days of release.
regulation on guava with reduced and stabilized The control of the striped mealybug was rated as
mealybug numbers and stable percentage parasit- outstanding (Mani et al. 1990). F. virgata
ism. The mean temperature during one sampling appeared in severe form on guava in the poly-
period was signicantly correlated with percent- house in 2006 in Bangalore North. A mean of
age parasitism 2 months later, indicating that 146.38 mealybugs/plant was observed on 5th
temperature has a major impact on parasitism March 2006 which declined to 0.96/plant on 7th
efciency. The encyrtid Leptomastidea abnormis May 2006 following the release of C. montrouz-
was responsible for about 90 % of the parasitism ieri. Cryptolaemus montrouzieri was the only
observed; the remainder was by another encyrtid, natural enemy recorded on F. virgata during the
Leptomastix dactylopii (Tingle and Copland study period. No other natural enemy was
1988). recorded on F. virgata infesting the potted guava
plants in the polyhouse (Mani 2008).
According to Zimmerman (1948), F. virgata
40.4.3 Ferrisia virgata was rst recorded in the Hawaiian Islands in
1898 but was a widespread and common pest in
Prothiophos, either alone or with mineral oil the Islands long before this. It is no longer com-
(0.5 %), gave better control of F. virgata than did mon in the Hawaiian Islands as it has been con-
malathion (the standard insecticide), the differ- trolled by the coccinellids Cryptolaemus
ence being evident from 8 days after application. montrouzieri, Olla v-nigrum and Azya luteipes,
Harvest residues were negligible. Mineral oil together with the syrphid, Alloagrapta obliqua.
(1 %) alone was also more effective than mala- At the beginning of a local outbreak, severely
thion but it caused leaf scorch followed by defo- infested branches should be cut and burnt imme-
liation. An ant barrier was ineffective (Villiers diately (Schmutterer 1969).
and de Stander 1978).
Aenasius advena Comp. and Blepyrus insu-
laris (Cam.), S. coccivora, Mallada boninensis 40.4.4 Planococcus lilacinus
(Okamoto), Brumus suturalis (F.) and Spalgis
epeus (Westwood), C. sexmaculata were Cryptolaemus montrouzieri supplements the
recorded on F. virgata (Mani et al. 1990). local natural enemies Brumoides suturalis
Chrysopa lacciperda (Kimmis) and Chrysoperla (Fabricius), Scymnus coccivora, Spalgis epeus
carnea were observed on F. virgata and P. citri in Westwood in clearing the population of P. lilaci-
40 Fruit Crops: Guava 383
invadens Williams, a serious exotic pest of horticulture Mani M, Krishnamoorthy A (2001) Suppression of
and other crops. Insect Sci Appl 13(3):411416 Maconellicoccus hirsutus (Green) on guava. Insect
Jalaluddin SM, Sadakathulla S (1998) Effect of Atso oil Environ 6(4):152
emulsion sprays in the control of mealy bug Mani M, Krishnamoorthy A (2007) Recent trends in the
Maconellicoccus hirsutus infesting guava fruits. biological suppression of guava pests in India. Acta
Entomon 23(2):151152 Hortic 735:469481
Joubert CJ (1964) Mealybugs on vines. Bull Dept agric Mani M, Krishnamoorthy A, Singh SP (1990) The impact
South Africa 243: 20 p of the predator, Cryptolaemus montrouzieri Mulsant,
Khalid M, Shafee AS (1998) Five new species of on pesticide-resistant populations of the striped mealy-
Pseudococcidae (Homoptera: Coccoidea) from North bug, (Ckll.) on guava in India. Insect Sci Appl
East India. Indian J Syst Entomol 5:6573 11(2):167170
Krishnamoorthy A, Mani M (1988) Records of green Manjunath TM (1986) Recent outbreaks of mealybugs
lacewings preying on mealybugs in India. Curr Sci and their biological control in Resurgence of sucking
58:155156 pests. In: S Jayaraj (ed) Proceedings of National sym-
Liu TS, Chang DC (1984) Population uctuations and the posium. TNAU, Coimbatore, pp 249253
control of citrus mealy bug on guava plants [Chinese]. Marotta S, Harten A, Van Mahyoub MA (2001) Mealybugs
Chinese J Entomol 4(1):8795 found on agricultural crops in Yemen. Bollettino di
Mani M (1994) Effectiveness of the exotic parasitoid Zoologia Agraria e di Bachicoltura 33(3):233238
Leptomastix dactylopii in the control of Planococcus Mukhopadhyay AK (2005) Study on the biology of the
citri in guava orchards. J Entomol Res 18:351355 mealybug, Ferrisicoccus psidii. Ann Plant Protect Sci
Mani M (1995) Studies on the natural enemies of orien- 13(1):239240
tal mealybug, Planococcus lilacinus (Ckll.) Sarwar M (2006) Occurrence of insect pests on guava
(Homoptera: Pseudococcidae) in India. J Entomol (Psidium guajava) tree. Pakistan J Zool
Res 19(1):6170 38(3):197200
Mani M (2008) Polyhouse efcacy of Cryptolaemus mon- Tanwar RK, Jeyakumar VS (2010) Papaya mealybug and
trouzieri Mulsant for the suppression of Planococcus its management strategies. NCIPM Techn Bull 22, 26 p
citri (Risso) on grapes and Ferrisia virgata (Cockerell) Tingle CCD, Copland MJW (1988) Effects of temperature
on guava. J Insect Sci 21:202204 and host-plant on regulation of glasshouse mealybug
Mani M, Krishnamoorthy A (1989) Record of Blepyrus (Hemiptera: Pseudococcidae) populations by intro-
insularis (Cam.) on Ferrisia virgata (Ckll.) in India. duced parasitoids (Hymenoptera: Encyrtidae). Bull
Curr Sci 58:644 Entomol Res 78(1):135142
Mani M, Krishnamoorthy A (1990a) Natural suppression Ullah GMR, Parveen A (1993) Coccoid pests (scale
of mealybugs in guava orchards. Entomon insects and mealybugs) and their host-plants on
15(34):245247 Chittagong University campus - a checklist.
Mani M, Krishnamoorthy A (1990b) Predation of Mallada Bangladesh J Zool 21(1):181182
boninensis on Ferrisia virgata, Planococcus citri and P Ullah GMR, Alam MS, Das HR (1993) Some aspects of
lilacinus. J Biol Control 4:122123 biology of pineapple mealybug, Dysmicoccus brevipes
Mani M, Krishnamoorthy A (1991) Breeding of Blepyrus (Cockerell) (Homoptera: Pseudococcidae). Chittagong
insularis (Hym., Encyrtidae) on Ferrisia virgate University Studies, Science 17(1): 7781
(Hemip., Pseudococcidae). Entomon 16:275277 Villiers EA, De Stander GN (1978) Control of the striped
Mani M, Krishnamoorthy A (1992) Contact toxicity of mealybug Ferrisia virgata on guavas [Afrikaans].
different pesticides to the encyrtid parasitoids Aenasius Citrus Subtrop Fruit J 541:1617
advena and Blepyrus insularis of the striped mealybug Williams DJ (1989) The mealybug genus Rastrococcus
Ferrisia virgata. Trop. Pest Manag 38:386390 Ferris (Hemiptera: Pseudococcidae). Syst Entomol
Mani M, Krishnamoorthy A (1993) Bionomics and man- 14(4):433486
agement of the striped mealybug, Ferrisia virgata Williams DJ (2004) Mealybugs of southern Asia. The
(Ckll.) a world review. Agric Rev 14:2243 Natural History Museum, London, UK and Southdene
Mani M, Krishnamoorthy A (1998) Regulation of SDN. BHD, Kuala Lumpur, Malaysia, 896 p
Rastrococcus iceryoides (Green) on guava. Insect Zimmerman EC (1948) Homoptera: Sternorrhyncha.
Environ 4:71 Insects Hawaii 5:1464
Fruit Crops: Mango
41
M. Mani
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
41.2.2 Planococcoides robustus (Pitan et al. 2002). After its appearance at the
eastern border of Cote dIvoire in 1989, the
It was found infesting the roots of mango in the mango mealybug, R. invadens rapidly became a
Kolar district of Karnataka, India. The mealy- nationwide constraint in mango production. On
bugs were enclosed in a parchment-like covering farmlands, 100 % yield losses could be reached
produced by a symbiotic fungus, but in May the so that infested orchards or trees were destroyed
surface of fruits that touched the soil was covered by farmers (Hala et al. 2004). The mango mealy-
with mealybug in various stages in the life- bug species are serious pests on mangoes in
history. Ants were seen to carry nymphs, which South Africa and result in considerable nancial
were active in the soil at this time (Puttarudriah losses due to the downgrading of mango fruits
and Eswaramurthy 1976). (Lagadec et al. 2009).
It was accidentally introduced into the West In Mexico, M. hirsutus was observed on Tommy
African region in the early 1980s, has become a Atkins, Haden, Manila, Ataulfo, Keitt, and Kent
serious pest of mangoes in several African coun- cultivars. Presence of insects was observed on
tries (Vogele et al. 1991). The spread of R. terminal buds and fruits in trees, as well as sur-
invadens in Nigeria was limited to Lagos, Ogun rounding weeds (Rosas-Garcia and Parra-
and Oyo States of the humid south-west contigu- Bracamonte 2011).
ous with the Republic of Benin. The frequency
with which infested plants were being felled,
burnt or sprayed with synthetic chemicals sug- 41.3 Natural Enemies
gested that the presence of R. invadens caused a
degree of panic by growers (Ivbijaro et al. 1992). 41.3.1 Rastrococcus iceryoides
On mangoes in the eld in Togo, all instars of R.
invadens were seen to stretch out the abdomen Several natural enemies were recorded on R.
from the leaf surface at a right angle if the leaves iceryoides in Northern India. Anagyrus pseudo-
were exposed to bright sunlight. In the labora- cocci Girault, Gyranusoidea sp., Praleurocerus
tory, the degree of lifting was found to be related viridis Agarwal, Allotropa sp., Microterys avus
to temperature. The reaction started at about 34 (Howard), Dinocarsis sp., Metastenus concinnus
C and reached a maximum at 37 C. Temperatures Walker, Cybocephalus sp., Scymnus coccoivora
measured in the eld on leaves exposed to bright Ayyar, Monomorium oricola (Jerdon),
sunlight were 34.541.1 C, indicating that the Coccophagus sp. and Proctolaelaps sp. were
reaction of the mealybug to high temperature can recorded from Malihabad in UP. But A. pseudo-
be interpreted as a heat-regulating mechanism cocci was found to be an important parasitoid of
(Agricola 1993). In Benin cv. Quinte WAS R. iceryoides (Tandon and Lal 1978; Rawat and
heavily infested with R. invadens and the cv. Jackmola 1970; Shafee et al. 1975; Sinha et al.
Gouverneur was slightly infested. The pre- 1985). Up to 42 % parasitism was observed in
reproductive period of R. invadens on the heavily nature in UP (Tandon and Lal 1978). According
infested tree was shorter and total offspring pro- to Narasimham and Chacko (1988), parasitoids:
duction greater than on the uninfested tree. Plant Anagyrus sp. nr. dactylopii (How.), Anagyrus sp.
genotype had the importance on R. invadens size nr inopus Noyes and Hayat., Coccophagus sex-
and survival (Boavida and Neuenschwander vittatus Hayat, Coccophagus sp. (pseudococci
1995). Protein, fat, carbohydrate, ash, crude bre group), Praleuricerus viridis (Garwal), Allotropa
and moisture contents were depleted with sp., predators: Nephus sp., Leucopis sp.,
increase in mealybug population on mango plants Cacoxenus perspicax (Knab.), Spalgis epeus
41 Fruit Crops: Mango 389
Coccophagus sp., Anagyrus sp., Gyronusodea Both physical and chemical control procedures
tebygi Noyes, C. montrouzieri, Spalgis epeus practiced by farmers have been ineffective
Westwood, Psectra inigua Hagen were recorded against R. invadens. It was observed that the
on R. invadens in India. Anagyrus sp. and G. mealybug is closely related to a complex of natu-
tebygi are worth trying parasitoids for trials ral enemies as parasitoids Gyranusoidea tebigi
against R. invadens in Africa (Narasimham and and Anagyrus mangicola in African counties
Chacko 1988). In West Bengal, Chartocerus sp., (Hala et al. 2004).
Azotus sp. [Ablerus sp.] and G. tebygi were The parasitoid Gyranusoidea tebygi native of
recorded on R. invadens (Das and Sahoo 2005). India was introduced into Togo in Africa in
November 1987 and released as a biological con-
trol agent against the mealybug on mangoes.
41.4 Management Establishment, spread and effectiveness of the
encyrtid were very good, resulting in satisfactory
41.4.1 Planococcoides robustus control (Agricola et al. 1990). In Togo and Benin,
both G. tebygi and Anagyrus mangicola were
Disulfoton as granule was applied to the soil capable of eliminating the mealybug (Moore and
monthly for a year, the plants were watered Cross 1992). The model predicted that the addi-
weekly, and the affected plants which had suf- tion of Anagyrus sp. to a system already contain-
fered desiccation and leaf-fall, showed signs of ing G. tebygi would lead to little improvement in
revival in India (Puttarudriah and Eswaramurthy the suppression of R. invadens (Godfray and
1976). Waage 1991). G. tebygi was released in Benin,
390 M. Mani
Gabon, Ghana, Nigeria, Sierra Leone and Zaire. important than the effect of G. tebygi (Bokonon-
In Togo, this parasitoid was established in all Ganta and Neuenschwander 1995).
areas infested by R. invadens. In addition, it In Nigeria, G. tebygi was released in 1991. By
established itself without previous release in 1997 and 1998, G. tebygi was found to have
Congo and Cote d'Ivoire. A. mangicola was crossed all agro-ecological barriers to colonize
released in Benin, Gabon and Sierra Leone since the entire area of infestation nationwide. During
1991 and by mid-1993 was recovered from a few this period, the populations of R. invadens had
sites. It seemed locally established in southern greatly decreased from between 11.0 and 98.0
Benin (Neuenschwander et al. 1994). mealybugs per leaf in 1991 to between 0.0 and
The mealybug population's potential rate of 18.2 mealybugs per leaf in 1998 (Pitan et al.
increase ranged from 0.066/day to 0.078/day. 2000). The population density of G. tebygi was
The potential for increase of the parasitoid was found to be negatively but signicantly corre-
double that of its host. In southern Benin, the lated with mango mealybug population and posi-
population density of R. invadens decreased dur- tively correlated with mango fruit yield.
ing the rainy seasons and peaked during the dry Parasitism was highly correlated with mealybug
seasons. Mealybug eld sex ratios were extremely population and yield, and was considered a major
variable, and the impact of such variability on the factor in the control of the pest and the subse-
mealybugs potential rate of increase was anal- quent increase in mango fruit yield. Rainfall did
ysed. The populations of the exotic encyrtid G. not have a signicant impact on yield, mealybug
tebygi, introduced into Benin in 1988 for control population or sooty mould score (Pitan et al.
of the pest, were synchronized with the mealybug 2002). In Ibadan, Nigeria, signicantly higher
populations. The spatial patterns of parasitism numbers of infested trees and mealybug popula-
distribution in relation to the host population tion, and signicantly lower parasitism levels
density were either independent or directly were found on mango trees in exhaust areas,
density-dependent, both at the tree level and for compared with others. Pollution level was corre-
larger zones. In the two orchards studied, mealy- lated with mealybug population (positively) and
bug populations eventually collapsed and disap- parasitism (negatively) in 2000 and 2007.
peared. It is concluded that the biological control Whereas mealybug population gradually built up
of the mango mealybug by G. tebygi was achieved on hitherto clean trees where pollution sources
by non-equilibrium local dynamics, and should were relocated, parasitoid activity seemed to be
be evaluated in a metapopulation perspective enhanced by the relocation of smoke sources.
(Boavida and Neuenschwander 1995). The effectiveness and conservation of the para-
In Benin, within 3 years, G. tebygi had colo- sitoids may therefore depend on the air quality
nized the entire area of infestation, and was found around the infested trees (Pitan 2008).
on practically all infested mango trees as well as Mango mealybug R. invadens an exotic pest of
other infested host plants. The percentage of mango was achieved with the release of parasit-
infested mango trees declined from 31 % in 1989 oids in African countries. Most producers attrib-
to 17.5 % in 1991. Average mealybug densities uted the observed improvement of mango
declined steadily from 9.7 females/48 leaves in production to the success of biological control.
1989 to 6.4 females/48 leaves in 1991. In multi- Based on production estimates by producers, the
ple regression analyses, based on 23 meteoro- negative impact of the pest on plant production
logical, agronomic and plant variables, the and the positive impact of the introduced natural
duration of the parasitoid's presence proved to be enemy were demonstrated. Interviewed mango
the major factor. It inuenced mealybug popula- producers gained on average US$ 328 per year by
tion densities and sooty mould incidence, which the biological control programme. Extrapolated to
in turn, affected the production of new leaves. In all producers of Benin, a yearly gain of US$ 50
all analyses, the impact of rainfall, for example, million in mango production was estimated. The
on the sooty mould or the mealybug was less value of accrued benets was estimated at US$
41 Fruit Crops: Mango 391
531 million over a period of 20 years. The total 41.5.3 Ferrisia virgata
cost of the biological control of mango mealybug
was estimated at US$ 3.66 million, which included At times, fruits were found covered with F. vir-
initial costs in other African countries and the gata in Tamil Nadu and Karnataka. C. montrouz-
introduction of the natural enemy from India, ieri was suggested for the suppression of the
resulting in a benetcost ratio of 145:1 for bene- mealybug (Anonymous 1987).
ts in Benin alone (Bokonon-Ganta et al. 2002).
The mango mealybug species are serious pests
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(Coccoidea:Pseudococcidae). In: Proceedings of the nomic evaluation. Trop Pest Manage 37(4):379382
National consulation meeting on strategies for deploy- Walker A, Hoy M, Meyerdirk DE (2003) Papaya
ment and impact of the imported parasitoids of papaya Mealybug. University of Florida featured Creatures.
mealybug classical biological control of papaya http://creatures.ifas.u.edu/fruit/mealybugs/papa
mealybug (Paracoccus marginatus) in India, pp 18 Williams DJ (2004) Mealybugs of southern Asia. The
Sinha PK, Sayeed MZ, Dinesh DS (1985) A report on the Natural History Museum/BHD, London/Southdene
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Fruit Crops: Papaya
42
M. Mani, M. Kalyanasundaram, and C. Shivaraju
Table 42.1 List of mealybugs recorded on papaya in dif- Barthlemy (Ben-Dov 2008), Guatemala (Ben-
ferent regions of the world
Dov 2008), Haiti (CABI/EPPO 2000), and
Species Country Reference Guadaloupe (Ben-Dov 2008) in 1998; French
Dysmicoccus Brazil Culik et al. (2006) Guyana (Ben-Dov 2008), Guiana (Ben-Dov 2008),
grassii Cuba Angeles Martinez et al. Guadeloupe (Matile Ferrero and Etienne 1998),
(Leonardi) (2001)
Cuba (CABI/EPPO 2000), and Puerto Rico in
Dysmicoccus Austro- Ben-Dov (1994)
1999 (CABI/EPPO 2000); Barbados (CABI/EPPO
nesophilus oriental
Williams & and 2000); Belize (Ben-Dov 2010), the Cayman Islands
Watson Pacic (CABI/EPPO 2000), Costa Rica (Ben-Dov 2010),
region Cayman and Montserrat in 2000 (CABI/EPPO
Ferrisia virgata Ben-Dov (1994) 2000), Nether lands Antilles (CABI/EPPO 2000),
(Cockerell)
the Bahamas and Guam in 20022003 (Meyerdirk
Maconellicoccus Florida Anonymous (2003)
hirsutus (Green)
et al. 2004); Palau in 2003 (Anonymous 2003;
Niapecoccus Ben-Dov (1994) Muniappan et al. 2006) and neighbouring islands
nipae (Makell) in the Pacic (Meyerdirk et al. 2004); Hawaii-Maui
Planococcus and Oahu in 2004 (Heu and Fukada 2005; Heu
citri (Risso) et al. 2007), the Northern Marianas (Tinian) in
Planococcus Trinidad Francis et al. (2012) 2005, and the Northern Marianas (Tinian) in 2005.
minor (Maskell) In Africa, it was reported in Ghana in 2009
Phenacoccus India (Cham et al. 2011). It was noticed in South and
solenopsisTinsley
South East Asian region during 20082009. In
Pseudococcus India Mani et al. (2013a, b)
jackbeardsleyi May 2008, it was recorded in Java, Indonesia and
Many Ben-Dov et al.(2001)
Gimpel and countries spread to Bali and Sulawesi Islands (Muniappan
Miller et al. 2008; 2009). It was also reported in July
Paracoccus Ghana Cham et al. (2011) 2008 in Colombo and Gampaha districts in Sri
marginatus India Cham et al. (2011); Lanka (Galanihe et al. 2010), Joyedpur in
Williams and Shylesha et al.
Granara de (2011d); Tanwar et al.
Bangladesh; Phnom Penh in Cambodia in 2010,
Willink 2010; Mani Manila in Philippines in 2008; Thailand in 2010
Chellappan (2011a); (Muniappan et al. 2009). The pest was rst
Muniappan et al. reported from Negeri Sembilan and Selangor in
(2008); Jacob Mathew
(2011)
Malaysia in February, 2009 (Muniappan et al.
Florida Walker et al. (2003),
2008; Mastoi et al. 2011 ) and Taiwan in 2010
Miller and Miller (Chen et al. 2011) and Maldives very recently.
(2002) In India, it was found at Coimbatore in July
Sri Lanka Galanihe et al. (2010) 2008 in Tamil Nadu (Muniappan et al. 2008;
Malaysia Mastoi et al. (2011) Regupathy and Ayyasamy 2009; Suresh et al. 2010).
Puerto Pantoja et al. (2007) Since July 2008 from Coimbatore in Tamil Nadu, it
Rico has spread subsequently neighbouring states such
Indonesia Muniappan as Karnataka, Andhra Pradesh, Maharashtra,
et al.(2008)
Kerala, Tripura, Jorat and Orissa in India (Shylesha
Hawaii Ronald et al. (2007)
et al. 2011c; Rabindra 2010; Krishnamoorthy and
Palau Muniappan
et al.(2006) Mani 2011; Sajeev 2011; Jacob Mathew 2011;
Pseudococcus Ben-Dov et al.(2001) Mani Chellappan 2011a; Chandele et al. 2011; Lyla
viburni and Philip 2010; Krishnakumar and Rajan 2009;
(Signoret) Mahalingam et al. 2010; Suresh et al. 2010).
Pseudococcus Ben-Dov et al.(2001)
longispinus
(Targioni
Taxonomy Paracoccus marginatus Williams &
Tozzetti) Granara de Willink (Hemiptera: Pseudococcidae)
42 Fruit Crops: Papaya 397
specimens were collected rst in 1955 in Mexico, stem. While feeding, it injects a toxic substance
but it was described in 1992 from the specimen into the leaves resulting in curling, crinkling,
collected in neotropical region (Belize, Costa rosetting, twisting and general leaf distortion
Rica, Guatemala and Mexico) by Williams and (Miller et al. 1999; Walker et al. 2003; Heu and
Granara de Willink (1992) and re-described by Fukada 2005; Pantoja et al. 2007). Heavy
Miller and Miller (2002) and also Angeles mealybug infestations render fruit inedible. Due
Martinez and De Los Suris (2005). Miller and to the build-up of thick white waxy coating and
Miller (2002) gave a complete description of all sooty mould development on the honeydew
the stages of the papaya mealybug. excreted by mealybug, infested fruits get reduced
market value. Fruits may fail to develop normally
Damage Papaya mealybug infestations are typi- and may be unusually small. Such fruits eventu-
cally observed as clusters of cotton-like masses ally shrivel and drop (Tanwar et al. 2010; Heu
on the above-ground portion of plants. Paracoccus et al. 2007). Some economically important crops
marginatus damages various parts of the host such as papaya, mulberry, cotton, cassava, citrus,
plant including the leaves, stems, owers and sweet potato, peas and beans, okra, eggplant,
fruits. P. marginatus may show very similar guava and ornamentals such as hibiscus,
symptoms to pink hibiscus mealybug Jathropha, Allamanda, Acalypha were severely
Maconellicoccus hirsutus (Green) (Pollard damaged by P. marginatus (Miller and Miller
1999). The insect sucks the sap by inserting its 2002; Mccomie 2000; Meyerdirk et al. 2004;
stylets into the epidermis of the leaf, fruit and Shylesha et al. 2011b).
Damage by P. marginatus
Ecology Mealybug occurs throughout the year ably does not tolerate cold conditions (CAB
but is active in warm dry weather. Prolonged International 2001). Heavy rains caused mortal-
drought with scanty rainfall and less number of ity of PMB especially of the crawlers stage.
rainy days favour the faster multiplication
(Ayyasamy and Regupathy 2010). During the
rainy season, papaya mealybug populations 42.2.2 Host Plants
decreased drastically because heavy rain washed
the insects off the plants. However, mealybugs It is highly polyphagus insect pest that can dam-
sheltered within unopened leaves and other hid- age large number of tropical and subtropical
ing places survived and built up their numbers fruits, vegetables and ornamental plants (Miller
again during the warm, dry weather. The climatic and Miller 2002). According to Muniappan et al.
preferences of P. marginatus have been docu- (2008), it was known to infest plants belonging to
mented well, but its occurrence in countries 22 families from Asia. Galanihe et al. (2010)
located 30 C from the Equator suggest that prob- recorded more than 40 plant species in Sri Lanka
398 M. Mani et al.
compared to 55 plants species recorded in Florida prevent the movement from one state to other
(Walker et al. 2003). Paracoccus marginatus states within the country (Tanwar et al. 2010).
attacks over 60 species of plants including eld
crops, fruit trees ornamentals, weed and scrub 42.2.4.2 Cultural Control
vegetation in India (Shylesha et al. 2011b). Planting material free from mealybugs is to be
used. In the initial stages of appearance of mealy-
bug, collection and destruction of infested plant
42.2.3 Natural Enemies parts are to be carried out (Ayyasamy and
Regupathy 2010; Tanwar et al. 2010).
It has never gained status as pest in the native
home of Mexico, Central and North America 42.2.4.3 Chemical Control
probably due to presence of endemic natural Chemicals were used desperately when there was
enemy complex (Walker et al. 2003). The papaya outbreak of mealybugs, and other methods were
mealybug became pest when it invaded the not available immediately. A number of insecti-
Caribbean region mainly due to the absence of cides like monocrotophos, methyl demeton,
natural enemies. Spalgius epeus Westwood was dimethoate, acephate, methomyl, fenthion, imi-
the predominant natural enemy on papaya mealy- dacloprid, thiomethoxam, dichlorovos, quinal-
bug damaging several host plants in South India phos, profenophos, fenitrothion, carbaryl,
(Thangamalar et al. 2010). Cryptolaemus mon- chlorpyriphos, diazinon, malathion, buprofezin
trouzieri Mulsant a general predator of mealybug were used against papaya mealybug (Tanwar
was also recorded occasionally on papaya mealy- et al. 2010; Regupathy and Ayyasamy 2009;
bug in India and elsewhere. Parasitoids of P. mar- Mahalingam et al. 2010; Banu et al. 2010; Suresh
ginatus from Mexico and Caribbean are listed by et al. 2010). They give short-term control but
Schauff (2000). Four species of chalcidoid para- chemical control is difcult and requires repeated
sitoids and two predators were found attacking application of the insecticides (Tanwar et al.
PMB in Malaysia (Mastoi et al. 2011). 2010; Ayyasamy and Regupathy 2010;
A total of 22 natural enemies occurring either Galanihe et al. 2010). The chemicals were
naturally/introduced were reported on papaya recommended for the control of the mealybug
mealybug in different countries (Mani et al. until the biological control agents could be
2012; Table 42.2. introduced.
42.2.4.4 Biopesticides
42.2.4 Management Fish oil rosin soap, azadirachtin and white min-
eral oils were found partially effective against
Mealybugs are difcult to control because they papaya mealybug. The three fungal pathogens
live in protected areas such as cracks, crevices Verticillium lecanii (Zimm.), Beauria bessiana
and under the bark of their host plants. Most of (Bals.) and Metarhium anisopliae (Metsch.) were
the stages including eggs of mealybug are cov- known to cause 4050 % mortality of P. margin-
ered with waxy secretions that protect them. An atus (Banu et al. 2010).
integrated pest management (IPM) approach
involving cultural practices, legal, chemical and 42.2.4.5 Biological Control
biological control is advisable. Though several methods were available, excel-
lent control of mealybug was obtained with use
42.2.4.1 Legal of biocontrol agent throughout the World
Strict quarantine measures are needed to prevent (Meyerdirk 2000). In the case of PMB also, out-
the entry of mealybug infested planting materi- standing control was achieved with use of para-
als/fruits/owers from other countries. Domestic sitoids in several countries (Mani et al. 2012;
quarantine measures are to be strengthened to Shylesha et al. 2011c).
42 Fruit Crops: Papaya 399
Parasitoids of P. marginatus
Meyerdirk and Kauffman 2001). However, Insects, Bangalore during JulyOctober, 2010.
Acerophagus sp. emerged as the dominant para- After ascertaining the safety in quarantine, these
sitoid species in both Puerto Rico and the three parasitoids were distributed to different
Dominican Republic (Meyerdirk and Kauffman states in India. Acerophagus papaya has done
2001; Ramirez and Sez 2002; Walker et al. exceedingly well in Karnataka & Andhra Pradesh
2003; Arnold 2001; Kauffman et al. 2001b). ( Shylesha et al. 2011c; Krishanamoorthy et al.
2011; Qadri et al. 2011), Maharashtra (Pokharkar
42.2.4.11 Florida et al. 2011; Mundale and Nakat 2011; Chandele
Paracoccus marginatus was discovered in et al. 2011; Nakat et al. 2011), Tamil Nadu
Florida 1998. The USDA Animal and Plant (Kalyanasundaram et al. 2011; Jonathan et al.
Health Inspection Service (APHIS) and USDA 2011), Kerala (Mani challappan 2011b; Jacob
Agricultural Research Service (ARS) initiated Methew 2011; Sajeev 2011), Orissa (Shylesha
a classical biological control programme for et al. 2011a) and Tripura state (Agarwala 2011)
the papaya mealybug. Four genera of encyrtid state in India.
endoparasitoid wasps specic to the mealybug C. montrouzieri was found colonizing on P.
were collected in Mexico by USDA and ARS marginatus in India (Shylesha et al. (2011b),
researchers and Mexican cooperators as poten- Malaysia (Mastoi et al. 2011), Palau (Muniappan
tial biological control agents: Acerophagus et al. 2008), Hawaii (Ronald et al. 2007), Florida
papayae, Anagyrus loecki, Anagyrus californi- (Walker et al. 2003), Guam (Meyerdirk et al.
cus and Pseudaphycus sp. (USDA 1999, 2000; 2004) and British Virgin Island (CAB
Meyerdirk and Kauffman 2001). A fth col- International 2001) but proved ineffective in
lected species was later reared and identied as checking the mealybug populations.
Pseudleptomastix mexicana (Noyes and
Schauff 2003). The rst releases of these four 42.2.4.13 Caribbean Islands
parasitoids were made in Florida in October As an exotic introduction to the Caribbean
2000 (Walker et al. 2003) and again released in islands, there were good prospects for control of
2003 (Meyerdirk 2003). Although it is believed P. marginatus by hymenopteran parasitoids origi-
that these parasitoids are established in the nating from its area of origin in Central America
released areas, Acerophagus papayae had (Pollard 1999).
higher per cent parasitism than A. loecki and
there is no recovery of P. mexicana (Kaushalya 42.2.4.14 Malaysia
et al. 2008). Paracoccus marginatus was reported for the rst
time in Malaysia on papaya, cassava, eggplant,
42.2.4.12 India jatropha and hibiscus plants. Four species of
Paracoccus marginatus invaded India in 2008 chalcidoid parasitoids were observed parasitizing
and has become severe on several agricultural the PMB. Acerophagus papayae was the major
and horticultural crops. The potential economic parasitoid of PMB. Two common predators
loss due to this pest ranges from 60 to 80 % in namely Apertochrysa sp. and Cryptolaemus
papaya. The parasitoids Acerophagus papayae, montouzieri were also found feeding on PMB
Pseudleptomastix mexicana and Anagyrus loecki (Mastoi et al. 2011).
from USDA-APHIS Puerto Rico were shipped to
India. A total of 3,429 of A. papayae, 1,485 of P. 42.2.4.15 Taiwan
mexicana and 516 of A. loecki were received by P. marginatus was found damaging papaya in
National Bureau of Agriculturally Important Taiwan for the rst time in 2011. A. papayae was
404 M. Mani et al.
Both adults and larvae of C. montrouzieri the population of mealybugs. Hence the reduc-
were found feeding on all the stages of JMB both tion of the mealybugs was attributed mainly to
in the eld and laboratory. A single predatory the action of/by all the three predators particu-
larva had consumed 3.83 (24), 13.75 (1214), larly C. montrouzieri (Mani et al. 2013a).
68.88 (6173) and 172.50 (164179) mealybug Williams and Watson (1988) state There are no
nymphs of 10 days old during the development of records of actual damage but the species is
rst, second, third and fourth instar, respectively. polyphagous and, in the absence of suitable natu-
The larva of C. montrouzieri took 13.85 days to ral enemies, it could be injurious. No classical
complete its development on JMB. The predator biological control attempt has been made for the
took 29.30 days on JMB (Mani et al. 2013a). Jack Beardsley mealybug, and possibly it is kept
under control by the local natural enemies in the
invaded countries (Muniappan et al. 2011).
42.3.3 Biological Control Hence there is no need for any panic for the new
invasive P. jackbeardsleyi in India (Mani et al.
Among the natural enemies C. montrouzieri was 2013a).
found in large numbers followed by S. epeus and
M. boninensis. The results on impact of natural
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Fruit Crops: Pineapple
43
M. Mani
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
short section of each of 3 or 4 inner whorl leaves. low and by the collapsing of the water storage
It is characterized by streaks of pale green to yel- tissues within these streaks.
Physical barriers such as ant fences running par- residual effects. The thick, waxy coating on
allel to the eld periphery are partially successful mealybugs makes insecticide penetration dif-
in keeping ants out of the eld, and subsequently cult. Even the use of systemic insecticides is fre-
controlling mealybug populations. quently impractical for mealybug control.
Pineapple industry, however, still needs an alter-
native for diazinon that can be used on mature
43.6 Cultural Control fruit prior to fruit harvest.
Dipping the basal portion of the planting
Previously infested elds should be turned over material in methyl parathion @ 0.02 to 0.05 % or
and all crop residues removed and burnt. Crop monocrotophos @ 0.02 % as a prophylactic mea-
residues and grass roots left in the eld may har- sure and application of carbofuran 3G @ 15 to 17
bour mealybug populations until the new crop kgha-1 in affected elds or phorate 10G @ 1.75
has developed enough to support a mealybug kgaiha-1 at 100 DAP can effectively control
population. Field borders should be kept clean of pineapple mealybug (Anonymous 2007). It indi-
weeds and debris that may support mealybugs cated that the basal portion of the planting mate-
between plantings. Weeds also provide alterna- rial needed double prophylactic measures
tive food sources that maintain ant populations (phorate 10 G and neem cake ground application
between periods where mealybug infestations are at 100 DAP and 180 DAP respectively), and
small. A common cultural practice is to allow a three times manual weeding helps to protect from
eld to lie fallow for 612 months after post- mealybug infestation (Mandal 2007). According
ratoon knockdown. This period is referred to as to the Pineapple Technical, PNB Krishi
the inter cycle. Shortly before replanting, the Samachar, Punjab National Bank expressed their
eld is burnt to remove pineapple trash. views that BCR in pineapple cultivation may be
1.92 and invest rupee return (IRR) may be more
than 50 % (Anonymous 2007).
43.7 Chemical Control
cally against this mealybug did not propagate pseudococcina Comp., were imported from Brazil
readily on the Hawaiian form, nor was establish- in 19351936 and further stocks of the latter spe-
ment secured with a long list of general preda- cies from Venezuela and Colombia. It was found
tors, among which were about 12 species of that the H. pseudococcina from Brazil, which is a
Hyperaspis and 6 species of Scymnus, presum- bisexual race, would not propagate on the
ably well adapted to attack on this mealy bug. Hawaiian D. brevipes, but the stock from
Some minor degree of control was attributable to Venezuela, which reproduces parthenogeneti-
the establishment of the cecidomyiid predator, cally, was well adapted to it. Both of the above
Vincentodiplosis pseudococci (Felt), imported species became established (Carter 1937).
from Mexico in 1930, and to a few of the numer- Anagyrus ananatis Gahan (Hymenoptera:
ous coccinellids that had been imported as gen- Encyrtidae) is the most common solitary, endo-
eral mealybug feeders (Fullaway 1924, 1933; parasitoid of PPM in Hawaii. The parasitoid has
Swezey 1925; Carter 1935; Zimmerman 1948). provided partial control of PPM in association
Mealybug species Pseudococcus bromelias on with other natural enemies. Field parasitization
pineapple was kept down by C. montrouzieri in of PPM by A. ananatis in the presence of ants can
Hawaii (Fullaway 1922). The encyrtid be as high as 9.9 %. It was present in all pineap-
Euryrhopalus schwarzi (How.) (=pretiosa ple elds surveyed and parasitized ant-tended
Timb.) and the cecidomyiid Dicrodiplosis guate- mealybugs (Gonzalez et al. 1999). Because of its
malensis Felt, both imported in 1935 from host specicity, abundance, and persistence, A.
Guatemala, have been reported as established. ananatis was chosen as a candidate for an aug-
Two encyrtid parasites, Anagyrus coccidivorus mentative biological control project targeted
Dozier to A. ananatis Gahan and Hambletonia against PPM (Gonzlez-Hernndez et al. 1999).
Mass production of a desired biological agent 14.8 C, they had emergence rates comparable to
is crucial to the implementation of any augmen- the control after 8 weeks, which indicated high
tative biological control program. The ability to survival rates at that temperature.
store reared biological control agents provides an
opportunity to manufacture them during low
demand periods and utilize them during high 43.10 Florida
demand periods. It also permits synchronized
eld releases of natural enemies during the criti- Although D. brevipes was only of very minor sig-
cal stages of pest outbreaks. Anagyrus ananatis nicance on a few small pineapple plantings in
prepupal and pupal stages could be stored for Florida, stocks of Hambletonia pseudococcina
over 6 weeks at 15 C without affecting their Comp. were imported from Puerto Rico in 1943
eclosion rate. When immatures were stored at 1944. The 1943 releases of very small numbers
416 M. Mani
were unsuccessful, but 374 adults released at 43.15 Taiwan and Bonnin Islands
three sites in 1944 resulted in establishment at
Sebring, Florida. C. montrouzieri was imported for the control of
D. brevipes but was only partially successful in
Taiwan and Bonnin Islands (Sakimura 1935).
43.11 Puerto Rico
Ferris GF (1950) Atlas of the Scale Insects of North technology interface, on 2829 December 2007.
America, Series 5: vii-f-278 p Bidhan Chandra Krishi Viswavidyalaya, Kalyani,
Fullaway DT (1922) Insect problems of pineapple indus- Nadia, West Bengal, pp 116
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Fullaway DT (1924) Insect collecting in Panama. Hawaii and wilt in pineapple. J Plant Protect Sci 1(1):4043
Forest Agric 21:9498 McEwen FO, Beardlsey JW Jr, Hapai M, Su TH
Fullaway DT (1933) Division of entomology. Hawaii (1979) Laboratory test with candi date insecticides
Forest Agric 30:5559 for control of the big-headed ant, Pheidole mega-
Gonzlez HH, Pandey RR, Johnson MW (2005) cephala (Fabricius). Proc Hawaii Entomol Soc
Biological characteristics of adult Anagyrus ananatis 23(1):119123
Gahan (Hymenoptera: Encyrtidae), a parasitoid of Menezes EB, Suzuchi J, Batista LB, Ismael AJ (1977)
Dysmicoccus brevipes (Cockerell) (Hemiptera: The use of granular insecticides for the control of the
Pseudococcidae. Biol Control 35:93103 pineapple mealybug, Dysmicoccus brevipes
Gonzlez-Hernndez H, Reimer NJ, Johnson MW (1999) (Cockerell, 1893) (Homoptera: Pseudococcidae)
Survey of the natural enemies of the pineapple mealy- [Portuguese]. Anais da Sociedade Entomologica do
bugs (Homoptera: Pseudococcidae). Biocontrol Brasil 6(2):287294
44:4758 Rao VP, Ghani MA, Sankaran T, Mathur KC (1971) A
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Jahn GC (1995) Gray pineapple mealybugs, Dysmicoccus Swezey OH (1925) Records of introduction of benecial
neobrevipes Beardsley (Homoptera: Pseudococcidae), insects in the Hawaiian islands. Hawaii Planters Rec
inside closed pineapple blossom cups. Proc Hawaii 29:369376
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Mallamaire A (1954) Les Homopteres nuisibles aux plan- Natural History Museum/Southdene SDN/BHD,
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Mamet N (1949) An annotated cataloque of the coccoidea tropical South Pacic region, Part 2, The mealybugs
of Mauritius. Bull Mauritius Inst 3:181 (Pseudococcidae). CAB International, Willingford,
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Fruit Crops: Avocado
44
M. Mani
44.1 Species
For some reason they will not stay under the pression and compression are followed by expo-
bags. The parasites, if they should nd the mealy- sure to ethanol vapour under decompression. The
bugs, would not be able to result in their death response of Hass avocado (cv. Hass) to MSDD
rapidly enough to prevent the destruction of the treatment for control of long-tailed mealybug
initial growth of foliage which is so vital to the (Pseudococcus longispinus) was investigated.
development of the scion. The prevention by the The best treatment for the most resistant life stage
parasites of the spread of incipient infestations of (2nd/3rd instars) was 90-min MSDD treatment
mealybugs is sufcient for control as far as the with 371 mg L1 ethanol. Early and late season
avocado tree as a whole is concerned, but on the Hass avocados were subjected to MSDD treat-
newly foliaged scions of grafted trees, prevention ments (with 371 mg L1 ethanol), or in air (con-
or immediate control is desirable. It is necessary, trol). Following the treatments, early season fruit
therefore, to turn to insecticides for the answer to were ripened at 20 and 25 C. Half of the late sea-
this problem. The long-tailed mealybugs appear son fruit were ripened at either 20 or 25 C, and
to the most abundant in late winter and early the remainder were stored at 5.5 C for 6 weeks,
spring, their numbers decreasing as summer then ripened at 20 C. There were no signicant
approaches. This period happens to coincide with difference in quality and rot incidence between
the period of greatest activity in the grafting of non-treated controls and MSDD-treated fruit. The
avocado trees. main disorders found were stem-end and body
rots, vascular browning and esh greying for the
stored fruit. There were also no signicant differ-
44.5 Insecticide Needed ences in fruit respiration rate or ethylene produc-
tion. Thus, MSDD was shown to be a potentially
Since prevention of attacks by the mealybug dur- soft disinfestation treatment for surface pests of
ing the entire late winter early spring period is the avocado (Blumberg and van Driesche 2001).
desired goal, an insecticide with a prolonged
residual effect against the young mealybug crawl-
ers, likely to become established on the grafted References
trees, might logically be, expected to be the solu-
tion to the problem. The insecticide mixtures were Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
applied with a paint brush to the top of the avo-
Pseudococcidae and Putoidae) with data on geograph-
cado stumps and for 3 or 4 inches down the side. ical distribution, host plants, biology and economic
It is especially important to apply the insecticides importance. Intercept Limited, Andover, 686 p
thoroughly to the area around each scion, both on Blumberg D, Van Driesche RG (2001) Encapsulation
rates of three encyrtid parasitoids by three mealybug
top of the stump and down the side. Imidacloprid
species (Homoptera: Pseudococcidae) found com-
(Condor Forte 200 SL) applied to the foliage monly as pests in commercial greenhouses. Biol
was efcient in controlling P. longispinus in both Control 22:191199
locations during the 2004 season in Chile. Dunkelblum E, Zada A, Gross S, Fraistat P, Mendel Z
(2002) Sex pheromone and analogs of the citrus
Applications to the trunk were not efcient against
mealybug, Planococcus citri: synthesis and biological
long-tailed mealybugs, apparently due to the activity. Bulletin OILB/SROP 25(9):213220
reduced absorption and translocation in the trees Flanders SK (1940) Biological control of long tail mealy-
at both locations (Sazo et al. 2006). bug on citrus and avocado. Calif Citrogr 25:146
Flanders SE (1944) Control of long tailed mealybug on
avocados by hymenopterous parasites. J Econ Entomol
37:308309
44.6 Quarantine Treatment Sazo L, Pizarro E, Araya JE (2006) Effect of the form of
application of imidacloprid on control of the long-
tailed mealybug Pseudococcus lingispinus (Targioni
Metabolic stress disinfection and disinfestations
& Tozzetti) on avocado and its impact on Neoseiulus
(MSDD) is a potential quarantine treatment in californicus (McGregor) in Chile. Boletin de Sanidad
which a combination of cycles of rapid decom- Vegetal Plagas 32(4):483490
422 M. Mani
Swirski E, Wysoki M, Yizhar J, Gurevitz E, Greenberg S Wysoki M, Izhar I, Swirski E, Gurevitz A, Greenberg S
(1979) The honeydew moth (Cryptoblabes gnidiella) (1976) Damage, host [range] and susceptibility of avo-
and the long-tailed mealybug (Pseudococcus longispi- cado varieties to the long-tailed mealybug
nus) in avocado plantations [Hebrew]. Alon Hanotea (Pseudococcus longispinus Targ. Tozz.)[Hebrew].
34(2):8386 Special Publication, Agricultural Research
Swirski E, Izhar Y, Wysoki M, Gurevitz E, Greenberg S Organization, Israel 65:8190
(1980) Integrated control of the long-tailed mealybug, Wysoki M, Izhar Y, Swirski E, Gurevitz E, Greenberg S
Pseudococcus longispinus (Hom.: Pseudococcidae), (1977) Susceptibility of avocado varieties to the long-
in avocado plantations in Israel. Entomophaga tailed mealybug, Pseudococcus longispinus (Targioni
25(4):415426 Tozzetti) (Homoptera: Pseudococcidae), and a survey of
Walter Ebeling, Pence RJ (1948) Control of the green- its host plants in Israel. Phytoparasitica 5(3):140148
house thrips, long-tailed mealybugs and June Beetles Zulhendria F, Jamiesona LE, Pererab CO, McDonalda
on Avocados. California Avocado Society 1948 RM, Connollya PG, Quekb SY, Woolfa AB (2012)
Yearbook 33:9395 The effect of metabolic stress disinfection and disin-
Williams DJ (2004) Mealybugs of southern Asia. The festation (MSDD) on Hass avocado fruit physiology
Natural History Museum/Southdene SDN/BHD, and mortality of longtailed mealybug (Pseudococcus
London/Kuala Lumpur, 896 p longispinus). Postharv Biol Technol 64:138145
Fruit Crops: Banana
45
B. Padmanaban and M.M. Mustaffa
B. Padmanaban (*) M.M. Mustaffa The waxy coating of the mealybug creates prob-
ICAR-National Research Centre for Banana, lem in getting desired results on mealy bug
Tiruchirappalli 620 102, India mortality with to insecticides. Therefore, use of
e-mail: bpadmanabannrcb@gmail.com
References
Beardsley JW (1986) Taxonomic notes on Pseudococcus
elisae Borkhsenius, a mealybug new to the Hawaiian
fauna (Homoptera: Pseudococcidae). Proc Hawaiian
Entomol Soc 26:3134
Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
Pseudococcidae and Putoidae) with data on geograph-
Ferrisia infestation ical distribution, host plants, biology and economic
importance. Intercept Limited, Andover, 686 p
Francis MA, Kairo WTK, Roda AL, Oscar E, Liburd OE,
Polar P (2012) The passionvine mealybug,
Planococcus minor (Maskell) (Hemiptera:
Pseudococcidae), and its natural enemies in the cocoa
45.6 Root Mealybugs agroecosystem in Trinidad. Biol Control 60:290296
Galanihe LD, Jayasundera MUP, Vithana N,
Asselaarachchi GW, Watson N (2010) Occurrence,
Since suckers are suspected to be the possible distribution and control of papaya mealybug,
source of infestation, spread of this serious pest Paracoccus marginatus (Hemiptera: Pseudococcidae),
(Geococcus spp.) is to be checked by prevention an invasive alien pest in Sri Lanka. Trop Agric Res Ext
13(3):2010
of use of suckers from infested areas within and
Huang T, Chien HS (1969) Chemical control of banana
outside the state. Dipping of suckers in boiled mealybug (Dysmicoccus brevipes (Cockerell)) (VII)
water for 10 s helps to destroy the live stages of [Chinese]. Plant Prot Bull (Taiwan) 11(3):123132
mealybug adhered to the sucker. Soil drenching Jones DR (1994) Risks involved in the transfer of banana
and plantain germplasm. In: Jones DR (ed) The
with chlorpyriphos 20 EC @ 2.5 ml per litre of
improvement and testing of Musa: a global partner-
water in the root zone helps to reduce the root ship. Proceedings of the rst Global conference of the
mealybug population. Sodium silicate was the International Musa testing Program held at FHIA,
best in reducing mealybug population on the Honduras, 2730 April 1994, INIBAP, Montpellier,
France, pp 8596
roots. Drenching of 3 % neem seed kernel extract
Lockhart BEL (1994) Banana streak virus In: Plotez RC,
(NSKE) and Verticillium lecanii (Zimmerman) Gentmeyer GA, Nishijima WT, Rohrbadr KG, Ohr
(Econil 7 g/l) were also effective against root HD (eds) Compendium of tropical fruit diseases.
45 Fruit Crops: Banana 427
American Phytopathological Society Press, St. Paul, -National conference on Banana held at Tiruchirapalli,
Minnesota, USA, pp 1920 2528 October 2007, pp 1819
Matile-Ferreo D, Williams DJ (1995) Recent outbreaks of Vargas Calvo A, Cubillo Sanchez D (2010) Evaluation of
mealybugs on Plantain (Musa sp.) in Nigeria including two modalities of management of the pseudostem
a new record for Africa and a description of a new spe- after the harvest on the growth, production and health
cies of Planococcus Ferris (Homoptera: of banana plants (Musa AAA) [Portuguese].
Pseudococcidae). Bulletin de la Societe Entomologique Agronomia Costarricense 34(2):287297
de France 100:445449 Watson GW, Kubiriba J (2005) Identication of mealy-
Meyer JB, Kasdorf GGF, Nel LH, Pietersen G (2008) bugs (Hemiptera::Pseudococcidae) on banana and
Transmission of activated-episomal Banana streak plantain in Africa. African Entomol 13(1):3547
OL (badna) virus (BSLOV) to cv. banana (Musa Williams DJ (1988) The distribution of the Neotropical
sp.) by three mealybug species. Plant Dis mealybug Pseudococcus elisae Borchsenius in the
92(8):11581163 Pacic region and Southern Asia (Hem.-Hom.,
Saranya C (2008) Evaluation of biocontrol agents against Pseudococcidae). Entomologists Monthly Magazine
Citrus mealy bug, Plannococcus citri. M.Sc. thesis sub- 124:123124
mitted to Bharathidasan University, Tiruchirapalli, India Williams DJ (2004) Mealybugs of southern Asia. The
Smitha MS, Maicykutty PM (2007) Root mealybug a Natural History Museum, Southdene Sdn. Bhd., Kuala
new pest problem on banana in Kerala. Abstracts Lumpur, 896 p
Fruit Crops: Sapota
46
M. Mani
from samples collected on 15th January. The cor- collected on 3rd March revealed the presence of
relation and regression analysis indicated a the exotic encyrtid parasitoid, Leptomastix dacty-
highly signicant relationship (r = 0.958) between lopii How. The parasitoid was found to be active
P. Citri (Y) and the parasitoid C. perminutus. The up to the rst week of April, 1996. The mealybug
regression equation tted with the mealybug population of 112.41 observed on 11th March
population (Y) and the parasitoid (X1) was: had declined to 2.16 on 4th April. Statistical anal-
Y = 20.5092 + 0.3912 X1. Abiotic factors (except ysis revealed that the parasitoid, L. dactylopii had
the minimum temperature) did not have any sig- a highly signicant relationship with the popula-
nicant relationship with P. citri. The decline in tion of P. citri (r = 0.969) during March-April.
the mealybug population from 156.4 in January The regression equation tted with the mealybug
to 2.05 in February 1996 attributed due to the population (Y) and the parasitoid (XI) was:
activity of the parasitoid C. perminutus (Mani Y = 7.56555 + 0.7644 XI. The relationship of the
and Krishnamoorthy 1997). mealybug population with any of the weather
In yet another orchard at llHR Farm, Bangalore parameters was not signicant. Hence, the reduc-
North, Planococcus citri was observed in the rst tion in the mealybug population during March
week of March, 96 on sapota. A mean of 94.37 April may be due to the activity of L. dactylopii
mealy bugs per shoot was observed. The samples (Mani and Krishnamoorthy 1997).
46 Fruit Crops: Sapota 431
46.3.4 Planococcus lilacinus Hodges AC, Hodges GS, Wisler GC (2005) Exotic scale
insects (Hemiptera: coccoidea) and whiteies
(Hemiptera: Aleyrodidae) in Florida tropical fruits; an
Cryptolaemus montrouzieri is highly effective in example of vital role early detection in pest prevention
reducing the populations of P. lilacinus and also and management. Proc Fla State Hortic Soc
the above other mealybug species on sapota. 118:215217
Mani M, Krishnamoorthy A (1997) Suppression of
Planococcus citri (Risso). Pest Manage Hortic Ecosyst
3(1):4547
References Mani M, Krishnamoorthy A (2008) Field efcacy of
Australian ladybird beetle Cryptolaemus montrouzieri
Agounke D, Agricola U, Bokonon-Ganta HA (1988) Mulsant in the suppression of Maconellicoccus hirsu-
Rastrococcus invadens Williams (Hemiptera, tus (Green) on sapota. J Biol Control 22:471473
Pseudococcidae), a serious exotic pest of fruit trees Mani M, Krishnamoorthy A, Pattar GL (2004) Efcacy of
and other plants in West Africa. Bull Entomol Res Cryptolaemus montrouzieri Mulsant in the suppres-
78:695702 sion of Rastrococcus invadens Williams on sapota.
Dhara Jothi B, Tandon PL (1991) Present status of sapota J Biol Control 18:203204
pests in Karnataka. Paper presented at National semi- Williams DJ (2004) Mealybugs of southern Asia. The
nar optimization of productivity and utilization of Natural History Museum/Southdene SDN/BHD,
sapota held at Navasari, India, 18 October 1991 London/Kuala Lumpur, 896 p
Fruit Crops: Pomegranate
47
M. Mani
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Table 47.1 List of mealybugs recorded on pomegranate in different regions of the world
Mealybug species Region/Country Reference
Crisicoccus theobromae Malaysia Williams (2004)
Williams & Watson
Dysmicoccus grassii Neotropical region Ben-Dov (1994)
(Leonardi)
Dysmicoccus neobreipes Thailand Williams (2004)
Beardsley
Ferrisia consobrina (Ckll.) India Williams (2004); Mani and Krishnamoorthy (1996)
Ferrisia virgata (Ckll.) India Nayar et al. (1976); Karuppuchamy (1994)
Pakistan Williams (2004)
Heliococcus destructor Palaearctic region Ben-Dov (1994)
Borchsenius
Maconellicoccus hirsutus India Mani and Krishnamoorthy (1991)
(Green) Maldives Williams (2004)
Nipaecoccus viridis India Mani and Krishnamoorthy (1990)
(Newstead)
Paracoccus ferrisi Ezzat & Mexico Ben-Dov (1994)
McConnel
Paracoccus marginatus Sri Lanka Galanihe et al. (2010)
Williams and Granara de Florida Walker et al. (2003)
Willink
Peliococcus trsipinosus Kenya Ben-Dov (1994)
(James)
Phenacoccus solenopsis
Tinsley
Planococcus citri (Risso) Soviet Union Niyazov (1969)
Iran Bodenheimer (1944)
Palestine Rivnay (1945)
Israel Rivnay (1960)
USSR Niyazov (1969)
Egypt EL-Rahn et al. (1974)
India Mani and Krishnamoorthy (1991, 2000)
Florida Hodges et al. (2005)
Planococcus dorsopinosus Philippines, India, Williams (2004)
Ezzat & Mc Connel Thailand
Planococcus lilacinus India Mani and Krishnamoorthy (1990); Ananda (2007); Balikai
(Ckll.) (2000)
India Tanwar et al. (2010)
Pseudococcus comsocki Ben-Dov (1994)
(Kuwana)
Pseudococcus cryptus India Williams (2004)
Hempel
Pseudococcus lamentosus France Frappa (1931)
Ckll.
Pseudococcus longispinus Africa Moawad et al. (2011); Wohlfarter et al. (2010)
(Targioni Tozzetti), Ps.
maritimus & Pl. citri
Pseudococcus comstocki Uzbekistan Sheffer (1974)
(Kuw.)
Pseudococcus maritimus http://ucce.ucdavis.edu/les/datastore/391-534.pdf
(Ehrhorn)
47 Fruit Crops: Pomegranate 435
In the Soviet Union, the main parasitoid of P. 47.5 Use of Verticillium lecanii
citri is Anagyrus pseudococci (Gir.), which
occurs in the south of European Russia and in Verticillium lecanii is known to infect the mealy-
Soviet Central Asia and which destroys up to 75 bugs infesting pomegranate in India. It is a cos-
% of the coccid population in areas not treated mopolitan fungus on insects. V. lecanii is known
with insecticides (Niyazov 1969). as white- halo fungus because of the white
mycelial growth on the edges of infested insects.
The conidia (spores) of V. lecanii are slimy and
47.4.3 Maconellicoccus hirsutus attach to the cuticle of insects. The fungus infects
insects by producing hyphae from germinating
Releases of the predator C. montrouzieri were spores that penetrate insect integument, and the
found to be very effective in controlling M. hirsu- fungus then destroys the internal contents and
tus on pomegranate in India (Mani and insects die. Treatments were imposed after the
Krishnamoorthy 1991). initiation of sufcient infestation of mealybug in
nymphal stage. It is seen from Table 4.1 that, all
the doses of V. lecanii i.e. 26 g/l of water effec-
47.4.4 Ferrisia virgata tively checked the built-up of mealybug popula-
tion up to 10 days after application and these
Scymnus coccivora and C. montrouzieri were treatments were found signicantly superior over
found to reduce the population of F. virgata in the untreated control at 3,7 and 10 days after
Tamil Nadu, India (Karuppuchamy 1994). application. All the V. lecanii treatments could
not cause mortality of the pest at 2 days after
application. The pest mortality was less (22.61
47.4.5 Pseudococcus maritimus 43.28 %) at 3 days after application than that
(39.3284.28 %) was observed at 7 days after
Mealybug biocontrol consists mainly of two application. It indicated that V. lecanii acts slow
kinds of parasitoids on Pseudococcus maritimus. initially and required at least 3 days to cause
The encyrtid parasitoids that help control mealy- lethal effect. On the basis of effectiveness of V.
bugs in grapes are also effective in pomegran- lecanii 4 g/l of water seemed to be optimum for
ates. Ladybird beetles with larvae similar in the effective management of mealybugs on
appearance to Cryptolaemus have been observed. pomegranate (Kulkarni et al. 2007). Effectiveness
The smaller encyrtid parasitoids rst appear of V. lecanii conidia and ltrates against
under the bark in the rst mealybug generation. Planococcus citri in vitro was reported by
There are probably ve parasitoid generations for Gonzalez et al. (1995).
each mealybug generation, as in grapes. The last
parasitoid generation occurs in mealybugs which
have already deposited half of an egg mass. The References
second encyrtid generation begins by parasitizing
crawlers under the bark and on the leaves, includ- Ananda N (2007) Seasonal incidence and management of
sucking pests of pomegranate. M.Sc. thesis submitted
ing those protected by the Dictyna spider webs.
to the University of Agricultural Sciences, Dharwad,
The larger parasite typically attacks large mealy- 59 p
bugs under the bark, so it appears late in each Ananda N, Kotikal YK, Balikai RA (2009) Management
generation. Biological control is effective unless practices for major sucking pests of pomegranate.
Karnataka J Agric Sci 22:790795
disrupted by Lannate or by ants (Devin et al.
Balikai RA (2000) Status of pomegranate pests in
2006). Karnataka. Pest Manage Hortic Ecosyst 6(1):6566
438 M. Mani
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Fruit Crops: Ber
48
M. Mani
48.1 Species ber and quality of fruits (Butani 1973). The infes-
tation with M. hirsutus on the growing point has
Mealybugs are injurious to ber (Zizyphus mauri- led to the malformation of shoots and leaves at
tiana L.) in India, Egypt, and Jordan etc (Table Bijapur in India. On an average, there were 80.6
48.1). colonies per plant, each colony having 17.8 indi-
viduals. On an average, there were 80.6 colonies
per plant, each colony having 17.8 individuals.
48.2 Damage Similarly 15.4 egg masses covered with white
waxy mealy matter were observed per plant.
In recent years, mealybugs have become an Based on the market price of infested and healthy
increasing threat to the cultivation of ber in penin- fruits, there was a net monetary loss of Rs. 25,800/
sular India. Mealybugs become serious pests in ha accounting for 33.33 % loss due to mealybug
Maharashtra, Andhra Pradesh, Karnataka, Tamil infestation (Balikai and Bagali 2000). The orien-
Nadu and Gujarat in India. Severe infestation of tal mealybug Planococcus lilacinus appeared in
mealybugs and subsequent development of sooty serious form on ber in 1990 and 1991 in Bangalore
mould affect the growth and fruiting capacity of North (Mani and Krishnamoorthy 1996).
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
on P. lilacinus (Mani 1993). There was reduction Hall WJ (1921) The Hibiscus mealybug Pseudococcus
hibisci (Hemip.). Bull Soc En dEgypt Cairo,
in numbers of P. lilacinus from 45.40/shoot in
XIV:1729
December 1994 to 0.40/shoot in rst week of Hall WJ (1926) The Hibiscus mealybug, (Phenacoccus
January 1995 due to the predation of S. epius in hirsutus Green) in Egypt in 1925 with notes on the
Karnataka (Mani and Krishnamoorthy 1996). introduction of Cryptolaemus montrouzieri Muls.
Minist Agric Egypt Bull 7, 15 p
Mani M (1993) Studies on mealybug and their natural
enemies in ber orchards. J Biol Control 7:7580
References Mani M, Krishnamoorthy A (1996) Biological suppres-
sion of oriental mealybug Planococcus lilacinus on
Balikai RA, Bagali AN (2000) Population density of ber. Pest Manage Hortic Ecosyst 2:4950
mealybug, Maconellicoccus hirsutus (Green) on ber Mani M, Krishnamoorthy A, Pattar GL (2007) Suppression
(Zizyphus mauritiana) and economic losses. Agric Sci of pink mealybug (Maconellicoccus hirsutus) on ber
Dig 20(1):6263 (Zizyphus mauritiana). Indian J Agric Sci 77(2):135
Barnes HF (1935) Some new coccids causing gall Meyerdirk DE, Khasimuddin S, Bashir M (1988)
(Cecidomyidae). Bull Entomol Res 26:525530 Importation, colonisation and establishment of
Ben-Dov Y (1994) A systematic catalogue of the mealy- Anagyrus indicus (Hym.: Encyrtidae) on Nipaecoccus
bugs of the world (Insecta: Homoptera: Coccoidea: viridis (Horn.: Pesudococcidae) in Jordan.
Pseudococcidae and Putoidae) with data on geograph- Entomophaga 33:229237
ical distribution, host plants, biology and economic Patil P, Patil BV, Patil PV (1996) Insect pests of ber in
importance. Intercept Limited, Andover, 686 p North Karnataka. South Indian Hortic 44:113
Butani DK (1973) Insect pests of fruit crops and their con- Shah AM, Patel CB, Patel VJ (1981) Ber, a new host
trol. Pesticides 7:3335 record of mealybug Nipaecoccus vastator (Mask.) in
EL Haidari HS, Aziz FI, Whahab WA (1976) Activity of Gujarat. Indian J Entomol 43:453454
predators and parasites of the mealybug, Nipaecoccus Tandon PL, Verghese A (1987) New insect pests of certain
vastator (Maskell) in Iraq. Yearbook of pl. Prot Res fruit crops. Indian J Hortic 44:121122
Iraq for 197476, pp 4146 Williams DJ (2004) Mealybugs of southern Asia. The
Natural History Museum, Southdene Sdn. Bhd., Kaula
Lumpur, 896 p
Fruit Crops: Custard Apple
49
M. Mani
Mealybugs are highly injurious to custard apple Fruits are completely covered with mealy-
(Annona spp.) in India (Mani and Krishnamoorthy bugs. When the population explodes, the
1989). Murray (1982) reported high level of infes- mealybugs are seen on the trunk and leaves
tation with P. citri on custard apple trees in but rarely. They cover the entire fruit reducing
Australia. In Caribbean islands, both Annona squa- the market value. Severe mealybug infestation
mosa and A. muricata were found severely infested causes heavy economic losses (Mani and
with M. hirsutus (Kairo et al. 2000) (Table 49.1). Krishnamoorthy 1989 ).
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Phalsa (Grewis asiatica Linn.) is cultivated in nellid Cryptolaemus montrouzieri Musant and
certain pockets of northern and peninsular India. the lycaenid Spalgis epeus Westwood were found
The pink hibiscus mealybug, Maconellicoccus clearing the populations of the mealybugs on
hirsutus (Green), is known to occur on the leaves, phalsa in the eld (Mani and Krishnamoorthy
owers and fruits of phalsa in India. The cocci- 1996).
Reference
M. Mani (*)
Indian Institute of Horticultural Research, Mani M, Krishnamoorthy A (1996) Reccord of two insect
Bangalore 560089, India pests and their natural enemies on phalsa. J Insect Sci
e-mail: mmani1949@yahoo.co.in 9(2):182
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Mealybugs are injurious to Jackfruit (Artocarpus In India, severe infestation of the spherical
heterophyllus) in India, Vietnam, Bangladesh, mealybug was observed on shoots of jack fruit.
Malaysia, Sri Lanka, Tonga, Caroline islands, The mealybugs suck the sap, leading to drying of
Solomon islands etc (Table 52.1). shoots in severe cases; fruits were also covered
with mealybugs (Mani and Krishnamoorthy
1997).
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Table 52.2 Nipaecoccus viridis and its natural enemies on Jack fruit (Mani and Krishnamoorthy 1997)
No. of healthy mealybugs/shoot No. of natural enemies emerged/shoot (Mean S.D.)
Date of sampling (N. viridis) A. dactylopii C. perspicax
8-3-1996 24.96 3.18 16.46 2.52 0.42 0.14
18-3-1996 16.15 2.47 15.07 2.02 1.53 0.37
27-3-1996 0.10 0.02 0.58 0.23 0.22 0.03
SD standard deviation
Paracoccus marginatus The parasitoid Sri Lanka, Thailand, Vietnam; Palaearctic: China,
Acerophagus papayae Noyes could be used to Saudi Arabia (Ben-Dov 2013). Coccidohystrix
control Pa. marginatus on tomato in India as it insolita has been a serious pest of brinjal
proved to be highly effective against the above (Solanum melongena) in many parts of India
mealybug infesting other crops. including Bihar (Lall et al. 1976), West Bengal,
(Chaudhuri 1976) and Kerala (Gopinathan et al.
Pseudococcus viburni Two potential biological 1982). Economic damage by mealybugs on brin-
control agents, the predator Cryptolaemus mon- jal was reported in Pakistan (Arif et al. 2009).
trouzieri and the parasitoid Leptomastix epona,
were considered for the use in controlling the Damage Both nymphs and adult mealybugs suck
mealybug Ps. viburni infesting tomato in the sap from leaves and tender shoots. Heavy clus-
Netherlands (Schoen and Martin 1999; Germain tering of mealybug C. insolita usually is seen
et al. 2003). Biological control of Ps. viburni was under surface of leaves as a thick mat with waxy
undertaken in greenhouses in France using a par- secretion. They also excrete copious amount of
asitoid wasp from Chile (Kreiter et al. 2005). honey dew on which the fungus sooty mould grow.
Affected plants appear sick and black, resulting in
Planococcus citri Anagyrus pseudococci is reduced fruiting capacity. If the ower blooms are
potential parasitoid of Planococcus citri infesting attacked, the fruit set is affected. When the fruits
tomato (Shariful and Jahan 1993). are infested, they can be entirely covered with the
mealybug. The infestation may lead to fruit drop
or the fruits remain on the shoots in a dried and
53.2 Brinjal/Egg Plant/Aubergine shrivelled condition. It is also a notorious pest of
stored potato tubers. The stored tubers are found to
Mealybugs are injurious to brinjal in many coun- be infested during July and October, when sprout-
tries (Table 53.2). The brinjal mealybug C. insol- ing of the buds takes place. Coccidohystrix insolita
ita has been recorded in Afrotropical: Kenya, was active in September-March on brinjal but
Madagascar, Rodriques Island (Mauritius), South found during April-August on alternative wild
Africa, Tanzania, Zanzibar; Australasian: Western host plants such as Solanum nigrum and Solanum
Samoa; Oriental: Bangladesh, Burma xanthocarpum, and had a mean life-cycle of 15.06
(=Myanmar), India, Laos, Pakistan, Philippines, days in India (Lall et al. 1976).
458 A. Krishnamoorthy and M. Mani
Varietal Tolerance Five aubergine lines/variet- gave excellent control of C. insolita in Andhra
ies were screened for resistance to C. insolita. An Pradesh (Tirumala Rao and David 1958) and
accession PI-381272-2 was found to be resistant Karnataka (Krishnamoorthy and Mani 1996) in
to C. insolita (Lit et al. 1998). India.
53.4 Chow-Chow
Pa. marginatus on hot pepper [Capsicum](Gimpel and Miller, 1996) Ph. peruvianus on chilli peppers
(Photo by Gevork Arakelian)
53.9 General Management trol of caretaking ants that are important for the
of Mealybugs in Vegetables proper development of mealybugs. Without the
ants, the populations are small and they spread to
Plant protection measures are of limited effec- new areas and elds would be slow and free from
tiveness against mealybugs because of its habit of serious infestations of mealybugs. Therefore,
hiding in crevices and the waxy covering of its management of mealybugs often includes the
body. Mealybug control often involves the con- control of ant species (Table 53.4).
464 A. Krishnamoorthy and M. Mani
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Proceedings of the National consulation meeting on marginatus Williams and Granara de Willink
470 A. Krishnamoorthy and M. Mani
Mealybugs are injurious to tuber crops, mainly cassava, and 6 species of mealybugs known on
cassava (Manihot esculenta), and to some extent cassava in West Africa. Mealybugs are most inju-
to taro (Colocasia esculenta), yam (Dioscorea rious in South America. In the 1970s, the cassava
spp.), sweet potato (Ipomea batatas Lam.), tan- mealybug appeared and threatened to decimate
nia (Xanthosoma sagittifolium), elephant foot the African cassava industry (Greathead 1978).
yam (Amorphophallus paeoniifolius), yam bean An account of mealybugs attacking cassava in
(Pachyrrhizus erosus), and enset (Ensete Neotropics and Africa is given by Cox and
ventricosum). Williams (1981). Paracoccus marginatus
(Williams and Granara de Willink) invaded sev-
eral countries and caused severe damage to cas-
54.1 Cassava sava (tapioca), particularly in India (Shylesha
et al. 2011). Stictococcus vayssierei (Richard
54.1.1 Species [Homoptera: Stictococcidae]), wrongly called as
cassava root mealybug, is really cassava root scale
Mealybugs are highly injurious in South America, (http://www.cabi.org/iscbeta/datasheet/118988).
Africa, India, Hawaii, Philippines, and Thailand According to Parsa et al. (2012), a total of 24 spe-
(Table 54.1). According to Williams (1978), 10 cies of mealybugs are known to attack Manihot
species of mealybugs are known in the world on esculenta. A list of mealybug species reported on
cassava in different regions is given in table.
Among the mealybug species, Phenacoccus
M. Mani (*) manihoti, Phenacoccus herreni, and Paracoccus
Indian Institute of Horticultural Research, marginatus are reported to cause heavy loss to the
Bangalore 560089, India cassava industry.
e-mail: mmani1949@yahoo.co.in
M. Kalyanasundaram
Tamil Nadu Agricultural University,
54.1.2 Phenacoccus manihoti
Coimbatore 3, India
C.A. Jayaprakas E.R. Harish R.S. Sreerag
The cassava mealybug, Phenacoccus manihoti
Central Tuber crops research Institute,
Trivandrum 17, India (Hemiptera: Pseudococcidae), is one of the most
severe pests of cassava in the world. Phenacoccus
M. Nedunchezhiyan
Regional Centre of Central Tuber Crops Research manihoti, the neotropical species (South
Institute, Dumuduma, Bhubaneswar 751 019, India America), was accidentally introduced to Africa
in the early 1970s, and it has become naturalized (Matile-Ferrero 1978; Neuenschwander et al.
throughout sub-Saharan Africa. Phenacoccus 1991; Williams and Granara de Willink 1992;
manihoti, an oligophagous mealybug, is consid- Zeddies et al. 2001).
ered as the major pest on an international scale
1991). In the absence of its natural enemies and Duration of rainfall had a lower effect, and rain
other control measures, this damage can reduce lasting 50 min or less caused less than 10 %
yields by more than 80 % (Nwanze 1982). reduction (le Ru and Iziquel 1990a).
Phenacoccus manihoti spread in Zaire occurs
over an area of 560,0002 km, mainly in regions
54.1.4 Varietal Susceptibility with a dry season of at least 90 days. Within the
region, the pest occurs principally in areas hav-
No cassava cultivars are known to be fully ing low green-leaf biomass, toward the end of
resistant to P. manihoti (Calatayud and Le R the dry season. Mealybug populations reached
2006). However, the Incoza variety was the catastrophic levels mostly during prolonged
most tolerant, followed by the Moudouma and periods of drought. The exotic parasitoid A.
Zanaga varieties. Dikonda, Kataoli, 3 M8, and lopezi has spread over 130,0002 km in western
1 M20 were highly susceptible (Tertuliano Zaire and southern Shaba, where no further
et al. 1993). The resistant clone 70,453 slows mealybug outbreak has since been recorded
down the buildup of mealybug populations and (Hennessey et al. 1990). In Nigeria, predicted
tolerates well the attack by mealybugs in Zaire yield losses in wet years were small, because
(Anonymous 1985). rainfall suppressed the population of P. manihoti
directly and enhanced the ability of the plant to
compensate for the feeding damage. In contrast,
54.1.5 Ecology losses in dry years were higher because of the
direct negative effects of water stress on photo-
In Ibadan, population peaks of P. manihoti usu- synthesis, which were compounded by the much
ally occurred during the second half of the dry larger population of P. manihoti that developed.
season (JanuaryApril). Apoanagyrus lopezi In Nigeria, the introduced encyrtid parasitoid A.
was the only natural enemy that was found dur- lopezi is the most important factor controlling
ing the whole year, and also in high densities the population of P. manihoti in the dry season,
(Hammond et al. 1987). In Congo, temperature and rainfall, directly or possibly via diseases,
appeared to be the most inuential factor on the during the rainy season. A. lopezi regulates P.
development time and on the capacity for manihoti in Nigeria, despite the disruptive effect
increase, which was highest (0.214) at 30 C of rain-induced mortality, drought effects on
and 75 % RH. Variations in abundance appeared host abundance, and predation by native cocci-
to be related primarily to the thermal conditions nellid beetles (Gutierrez et al. 1988). Severely
prevailing during the outbreak. Early multipli- infested cassava plants were lost between 9 %
cation of mealybug, beginning in July, under the and 46 % during the dry season, compared to
inuence of low temperature, would thus occur the pest-free plants. At the beginning of the
slowly, each generation being distinct with rainy season, mobilization of reserves from
clearly dened intervals in between, whereas a storage roots for regrowth caused temporary
late outbreak occurring under the inuence of root yield losses of up to 75 % (Schulthess et al.
high temperatures would develop more dramati- 1991). The population of Ph. manihoti was
cally, with a rapid succession of generations (le extremely low in all areas of Bolivia, Brazil, and
Ru and Fabres 1987) 1985. It was found that Paraguay, but there was a period of increase
intensity of rainfall seemed to be the most effec- from August to December (Lohr et al. 1990). In
tive factor, causing about a 22 % reduction. the Congo, during the wet season, torrential rain
476 M. Mani et al.
kept the populations at a low level by washing of the P. manihoti was observed in the Congo in
the insects (especially the crawlers) off the 1987. The development of the epizootic
plants. In the dry season, populations built up appeared to be more closely related to the fre-
rapidly, and outbreaks occurred, but these were quency of rainfall than to total rainfall.
rapidly reduced by the corresponding increases Conditions were highly favorable when the air
in the populations of natural enemies and in humidity was consistently greater than 90 % for
interspecic competition for the food plants, at least 5 h per day (le Ru and Iziquel 1990a;
from which many of the leaves had fallen, as a le Ru 1986).
result of the outbreaks (Fabres 1981). In Zaire,
the most important factor favoring multiplica-
tion and intensifying injury to the plants was hot
dry weather and dry eroded soil. The most effec-
Table 54.2 List of natural enemies recorded on P. mani-
tive control measures are therefore any cultural
hoti, F.virgata, and Ph.solenopsis infesting cassava
ones that help to conserve soil moisture, such as
Species Country References
erosion control, elimination of brush res, and
Phenacoccus manihoti
use of organic material as mulch or incorporated
Hyperaspis Nigeria Umeh (1983)
into the soil (Ezumah and Knight 1978).
marmottani (Fairm.)
Hyperaspis Congo Kiyindou et al.
senegalensis (1990)
54.1.6 Natural Enemies hottentotta
(Mulsant)
Explorations for the natural enemies of P. mani- Hyperaspis Congo Reyd and le Ru
raynevali (French) (1992)
hoti within its native range in South America
Hyperaspis Malawi Borowka et al.
(Bolivia, Brazil, and Paraguay) revealed the aestimabilis (Mader) (1997)
presence of four hymenopterous parasitoids Hyperaspis pumila Nigeria Iheagwam
(Angyrus sp. nr. pullus, Parapyrus manihoti sp.nr., (Mulsant) (1981)
Apoanagyrus diversicornis (Howard) Hyperaspis onerata Zaire and Bennett and
(Epidinocarsis diversicornis), Apoanagyrus (Mulsant) Congo Greathead (1978)
(=Epidinocarsis) lopezi (DeSantis), twelve pred- Ceratochrysa antica Nigeria and Barnard and
(Wlk.) Angola Brooks (1984)
ators, and one entomopathogenic fungus
Chrysopa sp. Nigeria Iheagwam
(Table 54.2), out of which the parasitoid
(1981)
Apoanagyrus lopezi appeared to be one of the
Exochomus Central le Ru and
most promising (Lohr et al. 1990; Pijls and Van aviventris (Mader) Africa Makosso (2001)
Alphen 1996; Noyes 1984). Apoanagyrus lopezi Congo Kiyindou et al.
takes 18 days to complete one generation (Odebiyi 1990)
and Bokonon-Ganta 1986). Exochomus troberti Malawi Borowka et al.
(Mulsant) (1997)
Exochomus avipes Gabon Boussienguet
(Thunberg) (1986)
54.1.7 Pathogens
Exochomus Congo Fabres and
concavus (Fursch) Matile-Ferrero
The epizootiology of the entomogenous fungus (1980)
Neozygites fumosa (Speare) in the populations (continued)
54 Tuber Crops 477
Nigeria, Rwanda, Senegal, Togo, Zaire, and ited with preventing the malnutrition of millions of
Zambia during 19811984 for the biological con- Africans and may well be the most important
trol of Ph. manihoti on cassava. Later, they were example of classical biological control ever.
introduced into cassava elds in over 100 locations
throughout sub-Saharan Africa. Apoanagyrus
lopezi was released in Nigeria in November 1981. 54.1.10 Congo
The spread of the parasite in a large cassava-grow-
ing area of Nigeria was at 5170 km/year, and Three severe outbreaks of Ph. manihoti have
became established in 16 African countries. A occurred since 1976. Phenacoccus manihoti pop-
reduction in the number of mealybugs to below the ulations declined greatly in the second year after
injury level was observed in every zone colonized the release of A. lopezi. (Hennessey and Muaka
by A. lopezi. In those zones, mealybug populations 1987; Hennessey et al. 1990).
reached peak densities of 1020 per terminal cas-
sava shoot or less, as compared with more than
1500 per shoot before the introduction of the para- 54.1.11 Nigeria
site. The introduction of this parasitoid into Africa
in the 1980s reduced high infestations by 90 %, Apoanagyrus lopezi was imported in 1981 from
becoming a highly successful case of classical bio- Paraguay into Nigeria for the biological control
logical control. Apoanagyrus lopezi is an efcient of Ph. manihoti (Lema and Herren 1985). Within
biological control agent across several ecological 3 years, it dispersed over 200,000 km2 in south-
zones of the African cassava (Neuenschwander western Nigeria, occupying 70 %98 % of all
and Hammond 1988). The wasp has been effective elds (Herren et al 1987). The impact assessment
in bringing the mealybug under control and revealed that 89 % of all sampled cassava tips had
reduces yield loss by 2.5 t per hectare. The suc- no individuals of P. manihoti at all
cessful control of the cassava mealybug problems (Neuenschwander and Hammond 1988;
has raised cassava yields and turned cassava into a Hammond and Neuenschwander 1990).
cash crop that is now spreading throughout Africa.
Zeddies et al. (2001) calculated the total costs and
benets of this biological control program for 27 54.1.12 Gabon
African countries over a 40-year period (1974
2013) under different scenarios, such as transport, The exotic encyrtid parasitoid Apoanagyrus
loss of crop, and even the price of maize as a pos- lopezi was introduced in Gabon for the biological
sible substitute. Based on the total cost of biologi- control of the cassava mealybug Ph. manihoti in
cal control at US$ 47 million, the benets from 1986. The establishment of the parasitoid after
different scenarios range mainly from 199:1 (or introduction showed a speed of dispersal of
US$ 9.4 billion) to 430:1 (or US$ 202 billion) 70120 km/year (Boussienguet et al. 1991).
(Williams and Granara de Willink 1992). Each
dollar spent on the mealybug control project
brought returns worth at least US$150 to the 54.1.13 Ghana and Ivory Coast
farmer. The overall economic benet of control-
ling the mealybug has been estimated at between P. manihoti in Ghana and Ivory Coast. P. mani-
US$9 billion and US$20 billion. Pedigo (1999) hoti populations were signicantly lower where
commented that the tremendous success is cred- A. lopezi had been present. In the savanna zone,
54 Tuber Crops 479
Cassava stem infested with P. manihoti in Thailand. Apoanagyrus lopezi from Benin to Thailand
480 M. Mani et al.
A. lopezi (500) that once saved Africas cas- declined sharply with the onset of rains (Van
sava farmers was brought on a ight from Benin Driesche et al. 1990).
to Bangkok. The Department of Agriculture is
now raising and releasing a quarter of a million
wasps in Thailand. The rst ofcial release began 54.1.20 Varietal Resistance
in July in the countrys northeast Tony Bellotti.
Instead of 10 years, it takes only a year or so to In Colombia, six clones (CM 21772, SG 100
respond (Paul Cox 2010). 54, SG 2503, CM 60683, CM 52631, and SM
5408) were identied tolerant or moderately
resistant to P. herreni, which can cause yield
54.1.18 Phenacoccus herreni losses up to 88 % (Bellotti and Vargas 1991).
(Bellotti 1983). Preferential oviposition by Acerophagus coccois and Aenasius vexans were
Anagyrus diversicornis and Acerophagus coccois introduced from Venezuela. By the end of 1996, a
was second and third instar nymphs and adult total of 35,930 parasitoids had been released. In
mealybugs. Bahia, A. diversicornis was recovered at 130, 234,
304, and 550 km from its release site after 6, 14,
Predators Six species of coccinellids, including 21, and 33 months, respectively. Acerophagus
Hyperaspis notate and H. onerata, were discov- coccois was recovered at 180 km from its release
ered in cassava elds infested with P. herreni in site, 9 months after release. Aenasius vexans,
Colombia (Carrejo et al. 1991; Castillo and however, did not disperse at all, despite being
Bellotti 1990; Sullivan, et al.(1991). The syrphid consistently recovered at its release site. In
Ocyptamus sp. was frequently found consuming Pernambuco, 9010 parasitoids were released from
eggs of P. herreni (Castillo and Bellotti 1990). In October 1995 onward. Acerophagus coccois and
Colombia, the main predator was found to be Aenasius vexans were recovered up to 40 km from
Ocyptamus stenogaster (Will.) forming 68 % of the release sites after 3 and 5 months of their ini-
all natural enemies in 1981 and Kalodiplosis coc- tial releases, respectively (Bento et al. 1999). The
cidarum (Felt) (11.6 %) in 1982 (Bellotti 1983). impact studies conducted between 1994 and 1997
indicated that at least 85 % of the parasitoids
Fungi P herreni, in May 1994, at Cruz das found in those elds were composed of the
Almas, Bahia, Brazil, were found to be infected recently introduced species Apoanagyrus diversi-
with Neozygites fumosa (Delalibera et al. 1997). cornis, Aenasius vexans, and Acerophagus coc-
In Colombia, a pathogenic fungus, Cladosporium cois. Apoanagyrus diversicornis was found in all
sp., was observed on P. herreni, being most effec- elds during most of the experimental period,
tive at high host densities (Bellotti 1983). whereas Acerophagus coccois and Aenasius vex-
ans were only found in the elds where they had
been released. Apoanagyrus diversicornis out-
54.1.22 Management competed Aenasius vexans in Sao Goncalo, but
not Acerophagus coccois in Itaberaba. The con-
Three encyrtid parasitoids, Apoanagyrus diversi- certed action of the three introduced parasitoids
cornis (Howard), Aenasius vexans (Kerrich), and and the native natural enemies was sufciently
Acerophagus coccois (Smith), are used to control efcient to control P. herreni at low levels in the
the cassava mealybug P. herreni in South America elds (Bento et al. 2000).
(Calatayud et al. 2001; Dorn et al. 2003a). For
efcient eld application, it is suggested to release
A. vexans and A. coccois late in the morning, dur- 54.2 Paracoccus marginatus
ing its period of increasing activity (Dorn et al.
2003b). A multispecies (A. vexans and A. coccois) Paracoccus marginatus (PMB), a polyphagous
approach to biological control of P. herreni may pest, is native of Central America/ Mexico, infest-
yield best results. ing more than 60 species of plants invaded over
In six states in northeastern Brazil, the mealy- 50 countries. In India, it was rst reported on cas-
bug P. herreni causes considerable damage to cas- sava from Tamil Nadu during 2008 (Muniappan
sava. Several native natural enemy species were et al. 2008), infesting a wide list of agricultural
found associated with the pest in Brazil but did and horticultural crops, including cassava/tapioca.
not provide adequate control. Exotic encyrtid par- Though Paracoccus marginatus was reported on
asitoids were imported and released in elds in cassava in more than nine countries, only in India,
the states of Bahia and Pernambuco. Apoanagyrus particularly in Tamil Nadu and Kerala, the cas-
diversicornis was introduced from Colombia, and sava crop was found severely damaged.
482
Heavy clustering of mealybugs was seen observed in a pre-release survey in all three loca-
under the leaf surface, giving the appearance of a tions. An average of 6.08 % parasitism and
thick mat with waxy secretion. They excrete 11.51 % reduction in papaya mealybug was
copious amount of honeydew that attracts ants recorded. The mealybug population had declined
and helps in the development of black sooty uniformly, corresponding to the gradual increase
mould, which inhibits the plants ability to manu- in percent parasitism at 2nd, 3rd, 4th, and 5th
facture food. This pest infests all aerial parts of months in all three locations. The average popu-
the plant. Infestation at the initial stage is lation of papaya mealybug from the tapioca gar-
observed on the leaf, particularly on the ventral dens was eliminated up to 93.15 % at 6th month
surface and petiole, and later it spreads to stems corresponding to 76.33 % parasitism. Parasitism
and branches. Heavy infestation causes leaf- by A. papayae accounted for 80.8994.31 %. It is
shedding and yield loss. The mealybug infesta- concluded that the release of A. papayae at 200
tion varied from 50 to 90 % in cassava, resulting individuals per location alone is sufcient to
in a monetary loss of Rupees 220 crores in cas- eradicate the population of papaya mealybug,
sava alone in Tamil Nadu. rather than the application of chemicals (Sakthivel
2013). Similar control of the mealyugg was
achieved with the release of A.papayae in other
54.2.1 Management districts, namely, Trichy, Erode (Divya 2012),
and Karur (Vijay 2010) in Tamil Nadu, India.
A comprehensive integrated pest management Tapioca is a major tuber crop of Kerala, and its
practices, viz., early detection by timely monitor- yield was reduced considerably by the infestation
ing, removal and destruction of affected plants of P.marginatus. In Kerala, total area for tapioca
and weeds, conserving natural enemies like pre- cultivation is 75,000 ha, and production is 30 t/ha.
dacious coccinellid beetles, lepidopteran preda- The mealybug infestation affected the tapioca pro-
tor, Spalgis epeus, and need-based application of duction to a great extent. Due to the release of
insecticides were developed. Even after adoption Acerophagus papaya in 2011, tapioca crop was
of IPM, the population of papaya mealybug was saved. Approximate cost of cultivation is Rs.
found to increase at a faster rate for want of ef- 50,000/ha, and the income is Rs. 3 lakhs/ha/year
cient natural enemies, since the pest is invasive (at Rs. 10,000/t). Thus, the net savings is 2.5 lakhs/
(introduced from other country), and the chemi- ha and 1.8 crores/year in Kerala, with the release of
cal control is short-lived and farmers have to Acerophagus papaya (Lyla, personal communica-
spray once in a fortnight. tion). Central Tuber Crop Research Institute,
Severe infestation of P.marginatus was Trivandrum (CTCRI) developed bioformulations
observed on cassava in Namakkal, Salem, and SHREYA and NANMA, which are very effec-
Dharmapuri districts of Tamil Nadu state tive against P.marginatus (http://www.yentha.com/
(Sakthivel and Qadri 2010), besides Coimbatore, news/view/4/bio-pesticides-developed-from-
Karur, Erode, Thirupur, and Trichy districts. tapioca-leaves).
Since Acerophagus papayae has provided excel-
lent control of P.marginatus in many countries, it
was imported from Puerto Rico in June 2010, and 54.3 Phenacoccus gossypii
releases were made in Tamil Nadu in 2010.
Population densities of papaya mealybug on Large populations of the mealybug Phenacoccus
cassava and percent parasitism in the three sam- gossypii (Tns. & Ckll.) had built up on cassava in
pling sites before and after release of parasitoids the Llanos Orientales, Colombia.. Seven species
are given in Table. Heavy population load of of parasitoids, of which the most important was
38.70, 43.85, and 41.21 numbers/5 cm2 was Anagyrus sp., and 18 species of predators were
recorded in Salem, Dharmapuri, and Namakkal observed in the eld. The most effective of the
districts, respectively. No parasitism was predators in controlling the populations of Ph.
484 M. Mani et al.
gossypii were Scymnus sp., Chrysopa sp., 54.5 Yam Dioscorea spp.
Coccidophilus sp., Ocyptamus stenogaster
(Will.), and Kalodiplosis coccidarum (Felt.) Three species of mealy bugs, including
(Milena Varela et al.1982). Planococcus lilacinus (Cockerell), P. citri, and P.
dioscorea have been reported to evoke a devastat-
ing impact on the yam tubers (Morse et al. 2000).
54.4 Sweet Potato Postharvest loss of tubers of yams and aroids due
to pests and diseases has ever been havoc. In
Paracoccus marginatus is known to attack a vari- warehouses and storage huts, insect pests are the
ety of crop plants, including sweet potatoes. The more serious menace to stored tubers and often
mealybugs feed, often in groups, on the underside more important than storage diseases. Damage
of leaves. Feeding on leaves by sucking out tissue inicted by the insects facilitates the entry of
uid, they release a poison into the plant tissue pathogens; besides, pests themselves indulge in
and secrete a sticky, sweet substance, called hon- spreading microbial contamination.
eydew. Where honeydew falls, a black fungal
growth called sooty mould develops. Although
this fungus does not harm plants directly, it blocks
out sunlight essential for the plant growth. The
mealybug infestations lead to stunted growth, dis-
coloration, malformed foliage, and defoliation of
sweet potato plants. To control papaya mealybugs
on sweet potato plants, gardeners should begin
with a biological approach by releasing natural
enemies, particularly Acerophagus papayae. For
severe infestations, applications at twice the nor-
mal rate of a pesticide, with an active ingredient
such as carbaryl or malathion, offer some control,
particularly when used alongside cultural mea- Mealybug damage to Dioscorea
sures (Tarah Damask What Causes White Bumps
on Sweet Potato Leaves? (http://homeguides.
sfgate.com/causes-white-bumps-sweet-potato- Feeding activity of the mealybugs not only
leaves-42692.html). Sweet potatoes are known to makes the tubers unattractive and unmarketable
be infested with mealybugs in storage. but also predisposes them to rot (Vasquez and
Buyser 2007; Rajamma et al. 2002). Infestation
of mealybugs leads to qualitative and quantitative
deterioration of the tubers, which culminates in
the unacceptability and low-prole marketability
of the tubers (Chomchalow, 2003). Palaniswami
and Pillai (1989) and Korada et al. (2010)
reported the qualitative and quantitative deterio-
ration of tubers of yams and aroids due to the
infestation by mealybugs. Palaniswami et al.
(1982) reported the incidence of Ferrisia virgata
(Cockerell) on sweet yam (Dioscorea dumeto-
rum). They desap the leaves, and the high inci-
dence of this pest causes drying of leaves and
Mealybugs on sweet potato withering.
54 Tuber Crops 485
54.6 Elephant Foot Yam the storage was for planting purpose, the corms
(Amorphophallus paeoniifolius) should be treated with fenitrothion (0.05 %) + man-
cozeb (0.2 %). (http://odisha.gov.in/e-magazine/
Amorphophallus paeoniifolius, popularly known Orissareview/2008/Sept-Octo-2008/eng-
as elephant foot yam, is an important tropical pdf/64-66.pdf). A two-instalment spray/drench
tuber crop in India. Mealybug (Rhizoecus amor- application of imidacloprid spaced 34 weeks
phophalli), a soft-bodied insect, infests the corms, apart resulted in complete control with zero phyto-
both in storage and in the eld (Misra et al. 2013). toxicity noted on any of the very mixed collection
Rhizoecus amorphophalli (Betrem) was widely (http://www.aroid.org/aroidl-archive/showthread.
distributed in South Asia, infesting many tuber php?id=3696). CTCRI-developed bioformula-
crops (Williams 2004). Mealybugs are seen in tions SHREYA and NANMA are very effec-
clusters on the stem, petiole, and leaf, particularly tive against the pests. Spraying neem oil at a
on the lower side. Infestation is high during warm concentration of 2 % at 15 days interval was found
and dry periods. Usually, the Amorphophallus effective to reduce the incidence (http://www.yen-
seed corms are harvested during the dry season, tha.com/news/view/4/bio-pesticides-developed-
after the crop is fully matured. During this period, from-tapioca-leaves).
mealybugs enter soil cracks and holes formed Though pesticides are effective in controlling
after drying of the pseudostem, and infest the mealybugs, they can be hazardous to human
corms. Infestation becomes severe when the health and the environment. Salt (NaCl) solution
corms are left for longer periods in the soil during (1000 ppm), cow urine, cow dung slurry (2 kg of
the dry season. Mealybug is a pest that thrives in cow dung in 1 L of water), and clay slurry (1 kg of
hot and humid conditions. When the temperature clay in 1 L of water) treatments were effective in
is more than 30 C, its infestation is severe, and reducing mealybug numbers and the associated
increases with rising temperature and humidity. corm damage. However, availability of cow urine,
Tubers are stored in storehouses, after harvesting, cow dung, and clay slurry limits their usage.
until further use. During storage, Rhizoecus amor- Common salt is cheap, widely available, and easy
phophalli (Betrem) causes 1015 % loss of tuber. to use in treating the corms prior to storage.
In the absence of mealybug control measures, Relative to untreated corms, those treated with
mealybug numbers increased by 45 times during salt solution recorded greater emergence when
the storage period. The pest affected the quality of eld-planted, as well as producing plants with
the corms and reduced subsequent eld establish- more vigorous growth (Nedunchezhiyan et al.
ment and crop growth. Infestation also affects the 2011). Cryptolaemus montruzieri was found pre-
corms ability to sprout, which then affects subse- dating on Rhizoecus amorphophalli in the storage.
quent production and productivity. Two species of It is recommended to maintain a temperature
mealybugs Ps.citriculus (Green) and Rhizoecus range between 25 C and 30 C in the elephant
sp. (Bit) are found together, and they suck and foot yam storage houses, as this temperature is
desap the cell contents of the tubers (Palaniswami most congenial for development, activity of
(1999). The eld infestation ranged from 6 to C. Montrouzieri, and for successful control of
45 %. Mealybugs multiplied during high tempera- mealybugs. Approximately two to three numbers
ture and humidity. They cover the tuber surface of C. montrouzieri are required for each infested
with powdery mealy substances. Severely infested tuber to control mealybugs. Accordingly, the
tubers shriveled, adversely affecting the quality predator can be released in storage godowns. As
and marketability. Several methods have been there is also natural parasitization of mealybugs
tried for controlling mealybugs. Rubbing of by Anomalicornia tenuicornis (Mercet)
infested corms with a dry cloth /soft brush, and (Encyrtidae, Hymenoptera), C. montrouzieri and
forcefully washing the corms with water are some A. tenuicornis together can contribute to the suc-
of the management practices which are cessful control of mealybug in storage, making
recommended. However, re-infestation after some the tubers suitable for planting during subsequent
time is common when using these techniques. If times (Misra et al. 2013).
486 M. Mani et al.
54.8 Enset the roots than the corms. Enset root mealybugs
were found up to a soil depth of 60 cm and up to
Presently, more than 12 million people in Ethiopia 80 cm from the corm. In addition, about 90 % of
depend on enset as a source of food. Its produc- the mealybugs were found within a 60 cm radius
tion is strongly hampered by the enset root from the plant (Addis, 2008).
mealybug Cataenococcus ensete (Williams and
Matile-Ferrero) in Ethiopia (Addis et al. 2008).
Enset plants infested with mealybugs have a 54.8.1 Management
retarded growth and dried lateral leaves. The
insects attack all plant age groups, but symptoms Repeated ploughing and sanitation of enset elds
are more severe on 2-to-4 years old enset plants. has also been reported as a control option for
Enset root mealybugs are found on roots and reducing enset root mealybug population num-
corms. Early infestations by root mealybugs can bers (Tadesse et al. 2003). Application of farm-
be easily overlooked, because they live under- yard manure (20 kg plant-1 year-1) resulted in
ground, and no visual symptoms will be observed vigorously growing plants with lower population
on the plant parts above the ground, until exten- numbers of enset root mealybugs (Anonymous
sive damage has been made to the roots and corm 2002). Among the insecticides tested, chloropyri-
(Hara et al. 2001). However, during periods of fos and diazinon have shown promising results
extreme drought, the mealybugs tend to move for its control and eradication (Tadesse 2006).
toward the corm when some of the roots drought. Soil drenching with diazinon 60 % EC and chlor-
The dispersal mechanism of enset root mealy- pyrifos 48 % EC caused at least 98 % mortality,
bugs is facilitated by the movement of infested both under eld and greenhouse conditions
suckers, farm implements during cultivation, (Tadesse et al. 2010). Still, cost-effective and
repeated transplanting operations, and associa- user-friendly control measures for the enset root
tion with ants. The population density of the mealybug have not yet been developed (Tadesse
mealybugs was signicantly (p < 0.05) higher on 2006) (Table 54.3).
Table 54.3 List of mealybugs recorded on tuber crops other than cassava
Crop and Mealybug
species Country Reference
Ipomoea batatas (Sweet potato)
Ferrisia virgata Guam IIse Schreiner (2000)
(Cockerell) Bangladesh http://www.aappbckv.org/journal/archive/6%20Sudden%
20outbreak% 20of% 20mealybug.pdf
Geococcus coffeae India David and Ananthakrishnan (2004)
(Green)
Maconellicoccus USA manatee.ifas.u.edu/comm-hort/pdf/pest-topics/InsectPHMHosts.pdf
hirsutus (Green)
Phenacoccus Ethiopia http://www.ppse.org.et/index.php?option=com_content&view=article
solenosis (Tinsley) &id=40:selonopsis-mealybug-phenacoccus-solenosis-tinsley-
hompotera-pseudococcidae-a-new-threat-to-cotton-production-in-
ethiopia&catid=7:-news&Itemid=3
(continued)
488 M. Mani et al.
Phenacoccus manihoti Mat.-Ferr. (Pseudococcidae) crops (Manihot esculenta) and observations on some
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Ornamental Plants
55
V. Sridhar, L.S. Vinesh, and M. Mani
Mealybugs are worldwide pests of ornamental citri (Risso), Phenacoccus solani Tinsley and
plants grown indoors and outdoors. Both green- Paracoccus marginatus Williams and Granara de
house and open cultivated grown ornamentals are Willink are some of the important mealybugs
commonly attacked by different mealybugs. In recorded on this crop.
recent years, mealy bugs have become an increasing
threat to the cultivation of several ornamentals in
India (Jhansi Rani 2001; Mani and Krishnamoorthy 55.1.1 Maconellicoccus hirsutus
2003). Mealybug infestation reduces vigour and
growth of the foliage which reduces the beauty of Hibiscus rosa-sinensis is a preferred and eco-
ornamental plant and affects marketability (Hamlen nomically important host of M. hirsutus also
1975). Mealybugs are a quarantine problem on popularly known as pink hibiscus mealybug
exported foliage and owers. Mealybugs cost grow- (PHMB), and is considered as a prolic pest that
ers and retailers millions of dollars per year in con- injects a toxin at the point of feeding, causing
trol costs and crop damage (Gullan and Kosztarab severe distortion of leaves and stunted growth
1997). An exhaustive list of mealybugs on various (Vitullo et al. 2009). Severe outbreak of M. hirsu-
ornamentals from different parts of the world has tus was noticed on ornamentals around Cairo in
been documented by Mattiuz et al. (2006), Cham 1920. Biological control is the best option for the
et al. (2011) and Arif et al. (2009) (Table 55.1). suppression of the pink hibiscus mealybug. The
parasitoid Anagyrus kamali Moursi and
Australian ladybird beetle, Cryptolaemus mon-
55.1 Hibiscus trouzieri Musant are the best-known natural ene-
mies to keep PHMB under check. Anagyrus
The greatest mealybug host diversity is found in kamali has been reported to be an outstanding
Hibiscus spp. Maconellicoccus hirsutus (Green), natural enemy in Egypt, Hawaii, Caribbean
Coccidohystrix insolita (Green), Planococcus islands and Florida, and is able to dramatically
suppress pink hibiscus mealybug populations.
The introduction of C. montrouzieri was facili-
V. Sridhar (*) L.S. Vinesh M. Mani
tated by India into Caribbean islands to control
Indian Institute of Horticultural Research,
Bangalore 560089, India M. hirsutus on several ornamental plants includ-
e-mail: vsridhar@iihr.ernet.in ing hibiscus (Gautam 2003).
Kuwana
Spilococcus leucopogi Ornamental plants Bulgaria Pencheva and Gerasimova (2006)
(Brittin)
Spilococcus eriogoni Asparagus California, Mexico Ben-Dov (1994)
(Ehrhorni)
Spilococcus Ornamentals Many countries Ben-Dov (1994)
mamillariae (Bouche)
Spilococcus pressus Nerium &Aster California Ben-Dov (1994)
Ferris
Vryburgia amaryllidis Plant families (Liliaceae and California & USA Lain et al. (2004)
(Bouche) Iridaceae)
Ornamental plants Bulgaria Pencheva and Gerasimova (2006)
Vryburgia bechuanae Geranium South Africa Ben-Dov (1994)
(Brain)
Vryburgia brevicruris Asclepiadaceae and other Belgium Ronse and Matile-Ferrero (1991)
(McKenzie) succulents
505
506 V. Sridhar et al.
Maconellicoccus hirsutus was known to attack ing levels. The population of C. montrouzieri
several ornamental crops in the Mariana Islands. declined for the rst 2 weeks, and then increased
The predator C. montrouzieri supplemented to a peak of 6 weeks after release. The predator
Anagyrus kamali and Allotropa mecrida sp. in population declined at about the same time as the
maintaining population density of M. hirsutus pest (McComie et al. 1997). In India, Spalgis
below the economic threshold at all locations epeus Westwood is the common natural enemy
(Reddy et al. 2009). After eld releases of C. found on mealybugs infesting hibiscus.
montrouzieri in May and July 1996 for control of Cryptolaemus montrouzieri, when released @ 20
M. hirsutus on ornamental hibiscus in Port of grubs/plant, reduced mealybug populations from
Spain, Trinidad, the mealybug population uctu- 84.3/plant in March to 0.9/plant in May (Mani
ated from 8 to 20 weeks, continuing, at a decreas- and Krishnamoorthy 2008).
According to Lai Yi-Chun and Chang Niann- sinensis (Goolsby et al. 2002). Since its acciden-
Tai (2007), all C. montrouzieri introduced were tal introduction into the island of Grenada in
killed and removed in 132.5 min by ants particu- 1994, A. kamali and C. montrouzieri were highly
larly Pheidole megacephala (Fabricius) and effective in bringing PHMB populations under
Tapinoma elanocephalum (Fabricius) in Taiwan. control (Sagarra and Peterkin 1999).
In Queensland, Australia C. montrouzieri was Invasion by M. hirsutus in Puerto Rico could
recovered on M. hirsutus infesting Hibiscus rosa- be restricted to less economic impact due to the
55 Ornamental Plants 507
timely introduction of A. kamali and of the laycaenid predator, S. epeus (Mani and
Gyranusoidea indica (Michaud and Evans 2000). Krishnamoorthy 2008). Cryptolaemus moun-
Efforts were made in 1922 to control M. hirsutus trouzieri was also found effective against this
on ornamentals with Cryptolaemus montrouzieri mealybug on hibiscus (Mani 2008).
introduced from France (Hall 1927). M. hirsutus
was found on ornamental hibiscus in Egypt in
20002001. Among several parasitoids recovered 55.1.4 Phenacoccus solenopsis
from PHMB, a gregarious parasitoid, Allotropa
mecrida was by far the most abundant parasitoid Phenacoccus solenopsis causes devastating dam-
attacking PHMB in Egypt. Primary parasitoids age on H. rosa-sinensis (Babasaheb 2012).
made up 94.9 % of the total parasitoids emerging Hibiscus syriacus was one of the most preferred
while 5.1 % were secondary (Gonzalez et al. hosts for P. solenopsis (Dhawan et al. 2010). A
2003). mean infestation of 96.4 % was reported in
Pakistan by Abbas et al. (2010).
According to Arve et al. (2011), the popula-
55.1.2 Planococcus citri tion of P. solenopsis on hibiscus was observed
throughout the year with its peak activity from
Planococcus citri causes severe damage to rst fortnight of October to rst fortnight of
Hibiscus rosasinensis. Cryptolaemus montrouz- December. The nymphs and adult female mealy-
ieri was effective for the management of this bugs were preyed by two predators Spalgis epeus
mealybug in India (Mani et al. 2011). (Westwood) and Scymnus coccivora (Ayyar
1963). Cryptolaemus montrouzieri gave excellent
control of P. solenopsis on Hibiscus after 4
55.1.3 Coccidohystrix insolita months of release in 2007 at Pune, India (Mani
2008). Aenasius bambawalei Hayat was found
Coccidohystrix insolita was recorded on H. rosa- parasitizing the Ph. solenopsis on hibiscus in
sinensis (Suresh and Mohanasundaram 1996 and Bangalore North. Hibiscus rosa-sinensis L. was
Williams 2004). Mealybug population declined found seriously infested with Phenacoccus sole-
from 145.6/plant in February to 0.6/plant in April nopsis in Guangzhou, Guangdong Province,
2003. There was 99.6 % reduction in the popula- China (Wu and Zhang 2009) and Iran
tion of C. insolita within 60 days of appearance (Moghaddam and Bagheri 2010).
was recorded in severe form on Jasminum sam- in India. Ferrisia virgata and Phenacoccus orna-
bac in India (Mani et al. 2011). Heavy popula- tus were also reported on Jasmine (David and
tions of Lankacoccus ornatus (Green) have been Ananthakrishnan 2004).
reported as covering the leaves of Jasminum sp.
55.13 Tube Rose August 2002. On 30th and 40th day of release,
the mealy bug population was further reduced
Mealybugs particularly Ferrisia virgata and to 50.45/plant and 4.87/plant, respectively. The
Planococcus citri have become increasing plants were completely cleared of the mealy
threat to tube rose cultivation in India (Mani bugs with about 50 days after the release of the
and Krishnamoorthy 2007a; Shanthi et al. predator (Mani and Krishnamoorthy 2007a).
2008). Following the release of C. montrouz- Dysmicoccus neobrevipes (Cockerell) is known
ieri, the mealybug population declined from to attack severely the roots of tube rose in
190 to 108/plant within 20 days during July Bangalore North, India.
leaf surfaces. It is advisable to check plants for dum (Linnaeus). Permanent establishment was
pests such as mealybugs before purchase and not achieved, however despite the fact that small
before introducing them into a greenhouse or colonies survived for some months (Bennett and
conservatory. Cultural control of bougainvillea Hughes 1959). Inspect the foliage of sago palm
mealybug can be achieved by removing and care- carefully to identify where mealybugs are located
fully disposing infested leaves and stems. The on the plant. Soak a cotton swab in rubbing alco-
larvae of Cryptolaemus are available to gardeners hol and rub the cotton swab over mealybugs on
and professional growers for the biological con- the infested palm to kill the bugs on contact. This
trol of mealybugs. They are most effective in treatment is effective for small numbers of mealy-
summer and need temperatures of at least 20 C bugs on a sago palm, but it is not practical to carry
for a few hours a day. For chemical control, sys- out when there are heavy infestations (http://
temic insecticides can be used. For ornamental www.ehow.com/how_12083815_kill-mealy-
plants in greenhouses or conservatories, garden- bugs-sago-palm.html).
ers can use products containing thiacloprid, acet-
amiprid, thiamethoxam or imidacloprid (http://
www.fera.defra.gov.uk/plants/publications/ doc- 55.26 Myoporum
uments/factsheets/bougainvillea.pdf).
In Morocco, C. montrouzieri was successful
some times against P. citri on the hedge plant
55.24 Araucaria (Myoporum) but was effective against heavy
infestations of P. citri when released in large
Araucaria, gliricidia and several other trees numbers (Boughelier 1935).
grown in parks and avenues in Bangalore (India)
have been found infested with the mealybug
Planococuus citri (Risso). C. montrouzieri was 55.27 MEALYBUG MANAGEMENT
the principal predator becoming abundant in IN ORNAMENTALS
April-May and October-November on these
plants. In some cases up to 12,000 larvae of Chemical Control
Cryptolaemus were encountered per tree. In such
cases, though belated, they completely controlled In Bulgaria, azinphos-methyl at 0.15 % and
the mealybugs (Manjunath 1986). phenthoate at 0.08 % proved the most effec-
tive materials against the mealybugs.
Pseudococcus maritimus (Ehrh.), Ps. adoni-
55.25 Sago Palm dum (L.) and Planococcus citri (Risso) infest-
ing ornamentals (Tsalev 1970).
Sago palms arent palm trees at allthese attrac- In Florida, acephate, oxydemeton-methyl and
tive, low-growing plants are actually cycads. Sago the insect growth regulator kinoprene were
palms are compact evergreen plants that make an found highly toxic to Pseudococcus longispi-
attractive addition to home interiors. The plants nus and Phenacoccus solani Ferris on orna-
are prone to mealybug infestations. Mealybugs mental plants. Overall reductions of Rhizoecus
are small, oval insects that use their sharp mouth- oridanus Hambleton by kinoprene and
parts to feed on the sap from sago palm foliage. epofenonane were comparable to the insecti-
Severe infestations can cause discoloured foliage, cides acephate and oxamyl (Hamlen 1977).
irregular growth and plant death. Mealybugs are One application of ZR-777 (prop-2-ynyl 3, 7,
easy to kill on sago palms with the proper treat- 11-trimethyl-(2E,4E)-2, 4 dodecadienoate) at
ment. In Bermuda, C. montrouzieri was liberated 1,000 ppm as foliar spray effectively reduces
on sage palm infested with Pseudococcus adoni- the mealybug populations on Ardisia crispa
516 V. Sridhar et al.
(crenata) and Gynura sarmentosa (Hamlen bugs. Insecticidal soap containing 49.5 %
1975). potassium salts of fatty acids or TweenReg con-
Diazinon, oxamyl, malathion, acephate and taining 100 % polysorbate 80 can be used to
butocarboxim all proved highly effective in create aqueous, plant-safe emulsions of these
controlling Planococcus citri on Dieffenbachia oils that are effective in controlling waxy insect
exotica, Ficus benjamina, Coleus sp. and Ps. longispinus. When sodium lauryl sulfate
Philodendron cordatum (Chandler 1980). and citric acid are included in the formulation,
This cocktail containing iminocloprid efcacy increases dramatically. Many types of
(growth regulator) + acephate (Systemic insec- ornamental plants tolerate these enhanced mix-
ticide) was sprayed on an orchid and coleus tures, which penetrate and kill mealybugs within
heavily infested with mealybugs. Two months seconds (Hollingsworth and Hamnett 2010).
later, these plants were ourishing with no Using 1 % limonene (a citrus extract), 0.75 %
sign of mealybugs. APSA-80 and 0.1 % Silwet L-77, a semitrans-
Acephate (780 mg/litre) was found to reduce parent mixture (primarily a micro emulsion)
the populations of longtailed mealybug was obtained that was safe for most plants and
Pseudococcus longispinus to zero on provided good control of mealybugs when
Asplenium bulbiferum (hen and chicken fern), sprayed or used in 1-min dips. Limonene has
while imidacloprid reduced the infestation to promise as a safe, natural pesticide for insect
14 % fronds infested and mean of 0.52.5 pests on tolerant plants. They caused no dam-
mealybugs on the youngest infested frond age to ornamentals with thick, waxy leaves,
(Martin and Workman 1999). such as palms, cycads, and orchids
In the Imperial Valley of California (USA), (Hollingsworth 2005).
imidacloprid-treated and thiamethoxam- The mealybug Phenacoccus solani Ferris was
treated hibiscus plants were completely free effectively reduced with acephate and vydate
of Maconellicoccus hirsutus (Castle and on Aphelandra squarrosa (Hamlen 1977).
Prabhaker 2011).
In Shanghai region 15 % aldicarb granules
mixed in soil around the ornamental succulent Biological Control
Kalanchos blossfeldiana plant roots or 40 %
omethoate 100x poured over the roots resulted Release the Australian ladybird beetle
in 97 % control or more of Planococus citri Cryptolaemus montrouzieri to control the
(Tang et al. 1992). mealy bugs on ornamentals in general (Mani
In Poland, pirimiphos-methyl 50 EC and and Krishnamoorthy 2003) and host specic
methidathion 40 EC are recommended to con- parasitoids for the control of mealy bugs on
trol Planococcus vovae (Nasonov) infesting ornamental crops.
common juniper, Juniperus communis (Golan In a commercial greenhouse in Leiden,
and Jaskiewicz 2002). Netherlands, successful control of the pseudo-
Horticultural oils kill all stages of the mealy- coccid Planococcus citri on Stephanotis
bugs that are present at application, and often plants was obtained with the coccinellid pred-
give good control. Oil products labelled as ators Cryptolaemus montrouzieri and Nephus
summer, superior, or Volck oil are high grade reunioni (Fursch); and the encyrtid parasitoids
and may be used on tolerant plants during Leptomastix dactylopii and Leptomastidea
either the growing or dormant seasons, but at abnormis (Girault) during summer and
different concentrations. autumn (Hennekam et al. 1987).
Post-harvest dips in properly formulated oil-in- Introduction of parasitoids Leptomastidea
water emulsions of terpene oils such as limo- abnormis and Leptomastix dactylopii gave
nene or essential oils such as peppermint oil or improved biological control of Planococcus
spearmint oil is advocated to control the mealy- citri in a large glasshouse stocked with a
55 Ornamental Plants 517
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Sciences and Veterinary Medicine Cluj-Napoca. Wysoki M, Izhar Y, Swirski E, Gurevitz E, Greenberg S
Horticulture 67(1):399402 (1977) Susceptibility of avocado varieties to the long-
Ullah GMR, Parveen A (1993) Coccoid pests (scale tailed mealybug, Pseudococcus longispinus (Targioni
insects and mealybugs) and their host-plants on Tozzetti) (Homoptera: Pseudococcidae), and a survey
Chittagong University campus a checklist. of its host plants in Israel. Phytoparasitica
Bangladesh J Zool 21(1):181182 5(3):140148
Vasundhara M, Siddappaji C, Kotikal YK (1990) New Yang JS, Sadof CS (1995) Variegation in Coleus blumei
record of downey snow line mealybug on Jasminum and the life history of citrus mealybug (Homoptera:
rigidum and its predators. Curr Res 19(6):99100 Pseudococcidae). Environ Entomol 24:16501655
Vennila S, Ramamurthy VV, Deshmukh A, Pinjarkar DB, Yang JS, Sadof CS (1997) Variation in the life history of
Agarwal M, Pagar PC, Prasad YG, Prabhakar M, the citrus mealybug parasitoid Leptomastix dactylopii
Kranthi KR, Bambawale OM (2012) A treatise on (Hymenoptera: Encyrtidae) on three varieties of
mealy bugs of Central Indian cotton production sys- Coleus blumei. Environ Entomol 26:978982
Orchids
56
N.K. Meena, R.P. Medhi, and M. Mani
ower buds, owers, and premature senescence. provides a substrate for the development of sooty
Such types of plants produce inferior quality ow- mould, which affects the rate of photosynthesis in
ers. In addition, mealybugs also produce honey- plants. Some species of mealybugs play a role as
dew, which makes the plant parts sticky, and vectors in the transmission of viral disease.
Mealybug infestation on Phaius ower bud Mealybug on leaf sheath of Phaius Mealybugon Cattleya leaf
All stages of the bug suck the sap from the plant temperatures, wind velocity, etc.) help to keep the
parts and secrete honeydew. Sooty mould develops population low or below economic injury level.
on the leaves in case of severe infestation. Attacked
plants look withered, reduced in growth with chlo-
rotic and deformed leaves. They prefer to live on 56.4 Mode of Mealybug Spread
roots deep in the media and are often only discov-
ered when orchids are repotted, though they will Orchids become infested with mealybugs in three
also attack other parts of the plant, especially under different ways: purchase of infested planting
the leaves. They will also hide in depressions on material, movement from infested to uninfested
pots, in sheaths, and in newly emerging growths plants, and windblown colonization. The occur-
(http://www.bellaonline.com/articles/art66287. rence of P. microcirculus is reported on the roots
asp). Imported exotic plants are inspected to pre- of orchids belonging to the genera Ansellia,
vent the accidental introduction of mealybugs in Cattleya, and Oncidium in the greenhouses in
Brazil (Camporese and Scaltriti 1991). The adult northern Italy (Lombardy). The orchids involved
females and nymphs of P. dendrobiorum are known had been recently imported from Brazil. With the
to infest leaves and roots of orchids in the green- entry of infested plants in an area where unin-
houses in the central part of the Korean Peninsula fested orchids are grown, juvenile mealybugs
(Kwon GiMyon et al. 2002). spread by crawling from one plant to another
plant through operational tools, irrigating water,
wind, pots, and potting media. Excess roots,
56.3 Seasonal Development dried leaves, pseudobulbs, and other wastage
thrown during repotting in the nearby areas are
The orchids are generally grown in greenhouses other most important modes of transportation of
or partially shaded net houses (under controlled mealybugs. Shifting of plants from one poly-
conditions), wherein mealybug species are active house to another polyhouse or from one place to
in warm climate (temperature range 2530 C), another place also plays a role in the spreading of
but in cold climate, when temperature goes below this pest. Sometimes, these mealybugs survive on
10 C, these mealybug species become less active unwanted plants (weeds), which provide suitable
and hide in protected places, such as among niche during unavailability of host or complete
roots, deep inside potting media, on pseudobulbs few stages of their life cycle during adverse con-
covering with scales, below leaf sheaths, and ditions and then migrate on the orchids. Ants that
other tight places. In open eld conditions, are attracted by honeydew produced by mealy-
mealybugs are susceptible to a variety of natural bugs can also be spread by crawling from one
enemies (predators and parasitoids), and weather plant to other plants.
factors (heavy rainfall, extremely high and low
56 Orchids 531
lay eggs. Unless the roots are checked and the application is less than botanical oils. Kinoprene
media changed, removal of mealybugs from only (tradename = Enstar II) is a synthetic form of
the upper plant portions is not a guarantee of suc- juvenile hormone, which is highly important in
cess. The potting medium can harbor eggs and insects at critical stages of their metamorphosis.
crawlers, so they should be disposed in a compost The use of kinoprene interrupts the normal devel-
pile or in the garbage. When repotting, a close opment of the insects, including mealybugs,
inspection, and, if necessary, a very gentle clean- scales, aphids, and whiteies. This insect hor-
ing and spraying of the roots before repotting are mone appears safe for humans and pets under
essential (https://www.aos.org/Default. usual use precautions. Experience on its use in
aspx?id=511). greenhouses and home collections suggest that
this may be the best new-generation pesticide for
controlling many orchid pests, including
56.8 Oils and Soaps mealybugs. Bifenthrin and other growth regula-
tors are also available for use on ornamentals, but
Horticultural oil, neem oil, mineral oil, and insec- little information is available for their use on
ticidal soaps are effective for mealybug suppres- orchids. Some of these new chemicals are very
sion. They are generally considered safe for effective, but are also highly regulated, and may
humans, pets, and plants than the usual insecti- not be available in some states for noncommer-
cides. None provide absolute control over mealy- cial uses. Azadirachtin (trade names = Azatin and
bugs, but frequent use during the presence of Neemazad) is a plant-derived chemical that is a
crawlers can serve to reduce their populations chitin inhibitor. Chitin is a primary compound
dramatically. The main caution with these oil used by insects when developing their integu-
solutions is that they should never be applied to ment or exoskeleton. Azadirachtin reduces the
plants on hot days (85 F) or in direct sunlight, as insects ability to properly develop its integument
to prevent burning of tissues (https://www.aos. and causes mortality through incomplete devel-
org/Default.aspx?id=511). opment. There is little information available on
this chemical for use on orchids, but more infor-
mation is available on its use on a wide variety of
56.9 Rubbing with Alcohol ornamentals, and is labeled for greenhouse appli-
cations, but may be too expensive for most home
As an orchid is a hard-leaved plant, gentle rub- greenhouse uses (https://www.aos.org/Default.
bing of leaves with cotton swabs dipped in aspx?id=511).
6070 % isopropyl alcohol gives satisfactory
results against mealybugs. If infestation occurred
in the root zone, inside potting media, the plants 56.11 Biological Control
should be removed immediately from the infested
media, and the roots should be sprayed with Use of natural enemies (parasitoids, predators,
chemical insecticides Further, repotting with and pathogens) for the management of mealy-
fresh media provides control measures to the pest bugs is the most effective and long-term solution
(https://www.aos.org/Default.aspx?id=511). in any crop ecosystem. Biological control agents
that are available commercially include a variety
of tiny parasitic wasps, brown lacewings, green
56.10 Growth Regulators lacewings, and lady beetles. The coccinellid bee-
and Chitin Inhibitors tles like Cheilomenes sexmaculata, Coccinella
septempunctata, and Cryptolaemun montrouzieri
These classes of insecticides have great potential are important predators of mealybugs (Lindemann
for use in orchid pest management. Growth regu- and Richter 2007). Cryptolaemun montrouzieri
lators are relatively expensive, but the cost per and Chrysoperla carnea larvae have been used as
56 Orchids 533
successful biological control agents of P. longis- (and money!), and such use allows resistant
pinus in potted Phalaenopsis orchids. mealybugs to develop. Finally, keep up the man-
Cryptolaemus montrouzieri was used to control ual removal of all mealybugs, if possible.
P. longispinus on the orchids in Germany Mealybugs are an excellent example of pests that
(Lindemann and Richter 2007). Pseudococcus are easily transported and that create tremendous
maritimus is becoming serious on the orchids in problems. Though most orchid keepers in North
India. Cryptolaemus montrouzieri can be tried America obtain their plants from conscientious
against all the arboreal mealybug species. growers in either Canada or the United States,
many persons do purchase plants while traveling,
in exchange from friends, or from questionable
56.12 General Management sources. Everyone needs to be aware of the great
Practices potential of inadvertently dispersing species to
new areas, particularly from international ori-
Heavy infestations of mealybugs, especially on gins. There cannot be enough stress placed on
many plants, require severe control methods the recommendation that all plants come from
using insecticides. On the extreme side, if a plant a reputable and quality grower, and are clean
shows signs of decline from infestation, then it is of pests.
seriously advised to destroy that plant, as the low
likelihood of rejuvenating that plant may not jus-
tify the expense and effort of continued treat-
ments. Also, destruction of a sick plant can be
References
used to justify the purchase of a new and health- Beardsley JW (1986) Taxonomic Notes on Pseudococcus
ier plant. If the mealybugs persist for long peri- elisae Borchsenius, a Mealybug New to the Hawaiian
ods of time (e.g., >9 months) even with the usage Fauna (Homoptera: Pseudococcidae). Proc Hawaiian
of the same insecticidal control method, then it Entomol Soc 26:3134
Ben-Dov Y (1994) A systematic catalogue of the mealy-
means probably that the mealybugs might have bugs of the world (Insecta: Homoptera: Coccoidea:
developed a resistant population. The best resolu- Pseudococcidae and Putoidae) with data on geograph-
tion to this is to change the methods and chemi- ical distribution, host plants, biology and economic
cals. The same chemical should not be used more importance. Intercept Limited, Andover, 686 p
Bronson CH (2009) An orchid mealybug, Pseudococcus
than three to four times sequentially. After isolat- dendrobiorum Williams (Hemiptera: Pseudococcidae).
ing the infested plants, it is suggested to give Pest Alert, Florida Department of Agriculture and
them a thorough application of something differ- Consumer Service, Florida, pp 13
ent from what has been used. For example, if Camporese P, Scaltriti GP (1991) Occurrence of
Pseudococcus microcirculus McKenzie (Homoptera:
insecticide is used, then switch on to an oil, soap, Coccoidea) on greenhouse cultivated orchids. Inf
or different insecticide. Resistance is not gener- Fitopatol 41(11):5961
ally a problem with growth regulators, such as Diaz A, Abreu N, Martin J, Suarez GM (2004) Hemiptera
kinoprene. Generally, an insecticide not labeled associated with wild orchids. Fitosanidad 8(2):4344
Gautam RD, Cooper B (2003) Insecticidal dip of some
for orchids should never be used. Whenever tropical cut owers for quarantine security against
using oils, soaps, and insecticides, be thorough, pink Hibiscus mealy bug, Maconellicoccus hirsutus.
change formulations frequently, and do not use Indian J Entomol 65(2):259263
less than the minimum concentration of mixture, Gimpel WF Jr, Miller DR (1996) Systematic analysis of
the mealybugs in the Pseudococcus aritimus complex
or more than that is normally recommended. Too (Homoptera: Pseudococcidae). Contrib Entomol Int
little of a chemical enhances resistance, while too 2(1):1163
high of a concentration may damage the plant. GiMyon K, Hwan LS, Jong HM, Gwan GH (2002) The
Unless you are a commercial grower rotating genus Pseudococcus (Westwood) (Sternorrhyncha:
Pseudococcidae) of Korea. J Asia Pac Entomol
mixtures of chemicals, do not use chemicals pro- 5(2):145154
phylactically, that is, do not routinely use chemi- Johnson PJ (2014) Mealybugs on Orchids. https://www.
cals as a preventative, as it is a waste of chemical aos.org/Default.aspx?id=511
534 N.K. Meena et al.
Jullien J (2009) What is your diagnosis? [French] Quel est Mandal D (2009) Eco-friendly management of
votre diagnostic? PHM Revue Horticole 510:4547 mealybug and wilt in pineapple. J Plant Prot Sci
Lindemann S, Richter E (2007) Biological control of 1(1):4043
Pseudococcus longispinus (Targioni Tozzetti) in Williams DJ (2004) Mealybugs of southern Asia. The
Phalaenopsis hybrids. Nachrichtenbl Dtsch Natural History Museum/Southdene SDN. BHD,
Panzenschutzdienst 59(4):7786 London/Kuala Lumpur, 896 p
Mahalakshmi V, Kalyanasundaram M, Karuppuchamy P, Yang SL (1997) Insect pests and harmful animals
Kannan M (2010) Biology and management of cotton of Phalaenopsis and their infestation habits.
mealybug, Phenacoccus solenopsis Tinsley Report of the Taiwan Sugur Research Institute,
(Hemiptera: Pseudococcidae). Entomon 35(2):7379 Taiwan, pp 4968
Medicinal Plants
57
V. Sridhar, L.S. Vinesh, and M. Mani
Mealybugs are injurious to several medicinal and farmyard manure (FYM) (12.5 t/ha) + Azophos (2
aromatic plants. Though medicinal and aromatic kg/ha) + neem cake (1000 kg/ha) and need-based
plants play an important role in public healthcare foliar application of neem oil (3 %) to be very
globally, they are affected by several mealybug effective in reducing the incidence of mealybug.
species. Incidence of P. solenopsis was observed Striped mealybug, Ferrisia virgata (Cockerell)
on a wide range of medicinal plants. Symptoms is another mealybug species, which causes dam-
of damage observed on these plants were twisted age on Aswagandha by sucking the sap from the
and dried leaves and shoot, white uffy mass on lower surfaces of leaves and pods during
stems, distorted or bushy shoots, presence of OctoberFebruary (Kumar 2007; Ramanna
honeydew, black sooty mould, small deformed 2009). Maximum population of 18 mealybugs
fruits, etc. (Chaudhary 2013). Various species of per plant was recorded during December 2008,
mealybugs recorded on medicinal plants and and the infested leaves turned yellowish and
cropwise options for their management are pre- dried up. Natural incidence of the predator
sented below. Cryptolaemus montrouzieri was observed on this
mealybug from Karnataka, India. Activity of
predators gradually increased from November
57.1 Aswagandha 2008 to January 2009 and then declined from
February 2009 onward (Ramanna 2009).
Coccidohystrix insolita (Green) (=Phenacoccus Solenopsis mealybug, Phenacoccus solenopsis
insolitus; Centrococcus insolitus) is one of the (Tinsley), was reported on Aswagandha from
key pests on Aswagandha (Withania somnifera) Tamil Nadu (Selvaraju and Sakthivel 2011).
(Williams 2004). Since Aswagandha is a herbal Abbas et al. (2010) reported a mean infestation of
medicine, application of synthetic chemicals 41 % by this mealybug on Aswagandha.
leads to accumulation of toxic residues. Hence, Papaya mealybug, Paracoccus marginatus
organic pest management including very safe (Williams Granara de Willink), an invasive pest
chemicals is the only option for this crop. was recorded in Tamil Nadu, India, in 2008, on
Ravikumar et al. (2008) found the application of papaya, and has attained the status of a serious
pest on a wide range of host plants, including
Aswagandha (Sakthivel et al. 2012; Selvaraju
V. Sridhar (*) L.S. Vinesh M. Mani
Indian Institute of Horticultural Research, and Sakthivel 2011).
Bangalore 560089, India
e-mail: vsridhar@iihr.ernet.in
Table 57.1 Various medicinal and aromatic plants infested with different mealybugs
Mealybugs Country/Medicinal plants, where recorded References
Paracoccus marginatus (Williams India Selvaraju and Sakthivel
Granara de Willink) (2011), Tanwar et al.
(2010)
Achyranthes aspera; Alternanthera sessilis:
Amaranthus viridis; Amaranthus spinosus;
Boerhavia diffusa; Calotropis gigantea; Cassia
angustifolia; Celosia argentea; Cleome
gynandra; Cleome viscose; Crotalaria retusa;
Glinus lotoides; Guettarda speciosa; Jatropha
gossypiifolia; Leucas aspera; Lippia nodiora;
Physalis minimam; Phyllanthus fraternus;
Phyllanthus amarus; Pulmonaria longifolia;
Solanum xanthocarpum; Tephrosia purpurea;
Trianthema portulacastrum; Tribulus terrestris;
Wedelia chinensis; Canthium inerme;
Phyllanthus niruri; Convolvulus arvensis;
Commelina benghalensis
India Chellappan et al. (2013)
Adhatoda vasica Nees, Alstonia scholaris (L.)
R. Br., Rauvola serpentina (L.),Benth. Ex.
Kurz, Cyanthillium cinereum (L.) H. Rob,
Bauhinia variegata, Ficus exasperate,
Azadirachta indica, Ocimum sanctum,
Couroupita guianensis, Indigofera tinctoria,
Cassia occidentalis, Phyllanthus amarus,
Phyllanthus fraternus, Datura stramonium,
Ghana Cham et al. (2011)
Wedelia trilobata; Sida sp.
Phenacoccus solenopsis (Tinsley) India Vennila et al. (2013), Vijay
and Suresh (2013a) and
Vijay and Suresh (2013b),
Nagrare et al. (2009), Saini
et al. (2009)
(continued)
57 Medicinal Plants 539
Selvaraju NG, Sakthivel N (2011) Host plants of papaya bug, Phenacoccus solenopsis Tinsley (Hemiptera:
mealybug (Paracoccus marginatus Williams and Pseudococcidoae). J Environ Biol 34:153158
Granara de Willink.) in Tamil Nadu. Karnataka J Agric Vijay S, Suresh S (2013a) Coccid pests of ower and
Sci 24(4):567569 medicinal crops in Tamil Nadu. Karnataka J Agric Sci
Singh MP, Ghosh SN (1970) Studies on Maconellicoccus 26:4653
hirsutus causing bunchy top in mesta. Indian J Sci Vijay S, Suresh S (2013b) Host plants of Phenacoccus
Ind 4:99105 spp. complex in Tamil Nadu. Karnataka J Agric Sci
Tanwar RK, Jeyakumar P, Vennila S (2010) Papaya 26:147151
mealybug and its management strategies, Technical Williams DJ (1986) The identity and distribution of
bulletin 22. National Centre for Integrated Pest Maconellicoccus Ezat (Hemiptera: Pseudococcidae)
Management, New Delhi, 22 p in Africa. Bull Entomol Res 76:621624
Varshney RK (1992) A check list of the scale insects and Williams DJ (2004) Mealybugs of south Asia. The Natural
mealybugs of South Asia. Part- 1. Rec Zool Surv India History Museum/Southdene SDN, BHD, London/
Occ Paper 139:1152 Kuala Lumpur, 896 p
Vennila S, Prasad YG, Prabhakar M, Agarwal M, Sreedevi
G, Bambawale OM (2013) Weed hosts of cotton mealy
Plantation Crops
58
Chandrika Mohan, P. Rajan,
and A. Josephrajkumar
The plantation crops viz., coconut, arecanut, Palmicultor palmarum, Phenacoccus sakai,
cocoa and tea are traditionally grown in India. Planococcoides anaboranae, Pseudococcus por-
The products and by-products of these crops tiludovici, Tylococcus malaccensis, Crinitococcus
form vital inputs for several industries and sus- palmae and Cyperia angolica are almost
tain livelihood for many million farm families. restricted to palms.
Infestation by insects form a crucial limiting
factor in attaining the production potential of
plantation crops. Among the sucking pest complex, 58.1 Coconut
mealy bugs, being polyphagous constitute a key
biotic stress that reduce plantation crop yield sig- In coconut, nine important species of mealybugs
nicantly. Fifty-seven species of Pseudococcids are reported from India viz. Palmicultor pal-
have been recorded on palms (Table 58.1). Half marum Ehron. Dysmicoccus cocotis Maskell,
of the Palmivorous species belong to the genera Pseudococcus longispinus Targ. Pseudococcus
Dysmicoccus, Planococcus, Pseudococcus and cryptus, Planococcus lilacinus, Pseudococcus
Rhizoecus. Among the wide array of mealy bug microadonidam, Nipaecoccus nipae Maskell,
species, Dysmicoccus has the most palmivorous Dysmicoccus nitimus and Rhizoecus sp. In
species (eight) including three species known Guam, Coccidohstrix insolita was observed
only from palms. The most commonly reported infesting coconut palm (Aubrey Moore et al. 2014).
mealy bug pest of palms are highly polyphagous Ferrisia virgata is also known to infest coconut
species, distributed worldwide and are primarily ( http://www.plantwise.org/KnowledgeBank/
known as pests of crops other than palms. Datasheet.aspx?dsid=23981).
Classical examples include Dysmicoccus brevi-
pes, Nipaecoccus nipae and Pseudococcus
longispinus. Few mealybug species namely 58.1.1 Palmicultor palmarum
Dysmicoccus hambletoni, Dysmicoccus cocotis,
Dysmicoccus nitimus, Neosimmondsia hirsuta, Palmicultor palmarum infests young seedlings
especially when they are closely spaced or in
nurseries and greenhouses. It does little damage
C. Mohan (*) P. Rajan A. Josephrajkumar to mature coconut palms but sometimes kills
ICAR-Central Plantation Crops Research Institute, seedlings. It has been observed in dense aggrega-
Kayamkulam, 690 533 Alappuzha district, tions on leaf axils especially on spindle/spear
Kerala, India leaves and at the base of spear leaves. Red ants
e-mail: cmcpcri@gmail.com
Table 58.1 Mealybugs recorded on Palms [Ben-Dov (1994), Howard et al. (2001)]
Mealy bug species Palm hosts Distribution
Chryseococcus arecae (Maskell) Rhopalostylis sapida Australia, New Zealand
Crisicoccus hirsutus (Newstead) Areca catechu India
Coccidohystrix insolita Green) Cocus nucifera Eastern Hemisphere, Guam
Crinitococcus palmae Ben-Dov Caryota sp. Philippines
Dysmicoccus boninsis (Kuwana) C. nucifera Pantropical
Dysmicoccus brevipes (Cockerell) Areca catechu, Carpentaria acuminata, Cosmopolitan (India, Indonesia,
C. nucifera, E. guineensis, Malaysia, Maldives, Philippines,
Pheonix dactylifera, Rhapis, Sri Lanka)
Roystonea & Sabal bermudiana
Dysmicoccus cocotis (Mask ell) C. nucifera Oceania, India
Dysmicoccus nitimus Williams C. nucifera Southern Asia, Malaysia
Dysmicoccus furcillosus sp.n. Areca catechu India
Dysmicoccus neobrevipes (Beardsley) C. nucifera Tropical America, Oceania,
Philippines
Dysmicoccus nesophilus Williams & Balaka seemanni Oceania
Watson
Dysmicoccus papuanicus Williams & C. nucifera New Guinea
Watson
Ferrisia consobrina Williams & Watson Metaxylon sagu Pantropical
Ferrisia virgata (Cockerell) C. nucifera & P. dactylifera Cosmopolitan
Formicoccus polysperes sp.n. Areca catechu India
Geococcus coffeae Green Chamaedorea Cosmopolitan
Laingiococcus painei (Laing) C. nucifera Oceania
Laminicoccus andersi Williams Gronophyllum & Howea Australia, New Zealand
Laminicoccus vitensis (Green ) C. nucifera & Roystonea regia Oceania
Leptococcus metroxyli Reyne C. nucifera & Metroxylon New Guinea
Maconellicoccus hirsutus (Green) P. dactylifera & P. sylvestris Cosmopolitan
Maculicoccus malaitensis (Cockrell) C. nucifera Oceania
Neosimmondsia esakii Takahashi Metroxylon amicarum & Ptychosperma Caroline Islands
ledermanniana
Neosimmondsia hirsuta Laing C. nucifera Solomon Islands
Nipaecoccus agathidis Williams C. nucifera Guadeloupe
Nipaecoccus nipae (Maskell) Areca sp. Arenga saccharifera, Cosmopolitan
Calyptrogyne, Chamaedorea,
Chamaerops excelsus, C. nucifera,
Gronophyllum, Howea belmoreana,
Howea forsteriana, Livistona chinensis,
Nypa fruticans, Pritchardia,
Ptychosperma, Rhapis humilis,
Sabal & Syagrus romanzofana
Nipaecoccus viridis (Newstead) C. nucifera India
Palmicultor palmarum (Beardsley) A. catechu, C. nucifera, Dypsis Oceania, Asia, tropical America,
lutescens, Latania glaucaphylla, Florida, Bermuda, Bangladesh,
R. regia & Veitchia sp. India, Malaysia, Maldives,
Philippines, Vietnam
Paraputo kukumi Williams C. nucifera Solomon Islands
(continued)
58 Plantation Crops 545
are mostly associated with this mealybug colony. 3789 offspring (Jalaluddin and Mohanasundaram
It was introduced into Florida where it has been 1993). Sometimes the red ant, Oecophylla sma-
observed in leaf axils and at the base of the spear ragdina (Fab.) was found associated with the
leaf of R. regia, Veitchia spp. and D. lutescens. In mealybugs.
severe cases, spear leaves become necrotic and
the palm dies. P. palmarum was found mainly on
coconut in Micronesia, Hawaii and Bahamas 58.1.2 Psuedococcus longispinus
(Williams 1981). P. palmarum has also been
recorded in Bangladesh on leaves of coconut and Pseudococcus longispinus affects spindle leaves,
palmyra palm (Borassus abellifer) (Ali 1987). and severe infestation results in failure of heart
In India, Palmicultor sp. took 21.60 days to com- leaf development and nally ends up with drying
plete its lifecycle. Adult females and males lived up of spindle. Seedlings are highly prone to
for 18.27 and 2.8 days, resp. A female produced attack.
58.1.8 Management the pest insects. The most important natural ene-
mies on coconut mealybugs are Pullus sp.,
Chemical Control Management of mealybugs Scymnus sp. (Coccinellidae), Spalgis epeus
begins with detection and identication of the (Lycaenidae), Bergineus maindroni
pest. Regular monitoring will allow detection of (Mycetophazidae), Dicrodiplosis sp.
these pests before damage is obvious and will (Cecidomyiidae), Homalotylus oculatus
also allow improved control. All plant parts need (Encyrtidae). These natural enemies exert good
to be searched, including the undersides of leaves control of the pest in nature. In case of severe
and stems. Pruning or washing infested plant infestation only, insecticides are to be applied. C.
parts can be helpful in reducing populations, par- montrouzieri was imported from California for
ticularly in cases of small infestations. A brisk the control of coconut mealybug Nipaecoccus
wash spray of water can also be helpful in reduc- nipae in Bermuda. The numbers released were
ing the population. Systemic insecticides can not probably adequate to provide reasonable
provide excellent options for mealybug control. opportunity for establishment (Bennett and
Soil application of thiodemeton [disulfoton] at Hughes 1959). In Seychelles, C. montrouzieri
0.5 g/plant and spraying with methyl demeton was introduced in 1959 and 1961 for the control
0.05 % were both highly effective against a of Pseudococcus (=Planococcus) longispinus but
severe attack of P. longispinus on 6-month-old not recovered (Bartlett 1977). The use of
coconut seedlings in India (Murthy and Pseudaphycus utilis Timberlake, a parasitic
Giridharan 1976). Application of 0.1 % mala- wasp, as a biological control agent successfully
thion, 0.025 % methomyl, 0.025 % demeton-O- controlled coconut mealybug Nipaecoccus nipae
methyl, 0.03 % dimethoate and 0.05 % in Hawaii and Puerto Rico (http://entnemdept.u.
phosphamidon caused 100 % mortality within 7 edu/creatures/orn/mealybug/coconut mealybug.
days, and 0. 05 % parathion-methyl caused 70 % htm).
mortality of Palmicultor sp. on coconut leaves
(Jalaluddin et al. 1991). Regular monitoring and
spot application twice with dimethoate 0.05 % at 58.2 Arecanut
20 days interval during summer to avoid further
spread of mealy bugs from infested coconut and Pseudococcus cryptus, Dysmicoccus brevipes
cocoa plantations (Nair 1983). and Dysmicoccus sp. are the mealybug species
found feeding on developing fruit bunches, spa-
dices, outer surface of leaf sheath, inorescence,
spindle leaves and occasionally on leaves. Severe
58.1.9 Biological Control infestation during tender nut stage causes imma-
ture nut fall.
Mealybugs are commonly attacked by predators,
parasitoids and diseases which can help manage
populations, particularly for long-term control. It 58.2.1 Pseudococcus cryptus
is important to recognize the presence of bene-
cial insects and to take steps to conserve them in It was found to infest leaves, inorescence and
the environment so they are available to control developing fruit bunches (Daniel 2003).
58 Plantation Crops 549
58.3.1 Planococcus lilacinus terminal buds, the ower cushions, the young
cherelles and the mature pods. Mealybugs feed-
The cocoa mealybug P. lilacinus (Ckll.) occurs in ing on the tender apical shoots result in reduced
most of the cocoa growing tracts of South East growth causing deformity of the shoots which
Asia viz., India, Sri Lanka and Papua New grow as brush-like structures. Infestation of the
Guinea. In India, it is reported as a serious pest ower cushion results in cushion abortion and
causing damage to cocoa and is present in all continuous attack results in withering and drying
cocoa tracts of the country. It is present through- of the ower cushions. The feeding on the bark of
out the year colonizing the tender parts of the pods by mealybugs results in irregular cracks and
plants such as the growing tips of the shoots, the pitting and feeding on cherelles result in cherelle
552 C. Mohan et al.
wilt. Peak population is reported in April-May example of NA34 and T63/967. Trees of T17/524
and low level of activity is recorded during rainy parentage were sparsely infested with mealybugs
and post monsoon seasons (Nair 1981). (Campbell 1990).
when it is hustled by the ant, aiming this one far the population of mealy bugs in cocoa plantations
from its safe place on the root (Delabie et al (Daniel 2002). In India, the predators observed
2008). with mealy bugs infesting cocoa are coccinellid
beetles, Scymnus sp., the lycaenid Spalgis epeus
Westwood. Trials with introduced predatory bee-
58.3.5 Management tles Cryptolaemus montrousieri did not give pos-
itive results (CPCRI 1986) probably due to
The control of mealy bug by insecticide is usu- activity of the associating attendant ant, mainly
ally difcult because of its habits, water repellent Oecophylla smargdina, in cocoa ecosystem.
nature of their body covering and the protection Ackonor and Mordjifa (1999) have listed the nat-
provided by the ant-constructed nests. Hence ural enemies of Planococcoides njalensis in
destruction of initial foci of infestation before Ghana. Natural enemies include two species of
attaining severe proportion is very important. coccinellid predators Hyperaspis sgregia Mader
Destruction of highly infested plant parts and and Scymnus sp., the cecidomyiid Coccodiplosis
removal of alternate weed hosts in the immediate coffeae Barnes and six hymenopteran parasitoids
vicinity aid in reducing the mealybug population. including Aenasius abengouroui (Risbec).
Locate ant colonies during summer ploughing Cecidomyiids predating on both species of
and destroy. Conservation of coccinellid lady Planococcus infesting cocoa was reported in
beetles in the ecosystem. Proper pruning of cocoa Southern Karnataka (Daniel 2002). Exotic para-
branches helps some way in preventing colony sitoid Acerophagous papayae was released to
build up of O. smaragdina (Daniel 2002). Nair control Paracoccus marginatus infesting cocoa
(1981) indicated that foliar spraying of 0.05 % inter-cropped in coconut garden at Kondikulam
fenthion, quinalphos or dimethoate was effective and Alivalam villages of Pattukottai taluk in
against P. lilacinus. This could even be applied as Thanjavur district in Tamil Nadu, India (http://
spot spray whenever the mealy bug population www.thehindu.com/todays-paper/tp-national/tp-
was over 15 % for maintianing the population at tamilnadu/parasitoids-to-control-mealy-bug- -
a lower level. infesting-cocoa-released/article1157926.ece). In
When the infestation is lesser, Spraying of the area where P. marginatus alone occurs, eld
neem oil 3 % or sh oil rosin soap 25 g/litre was release of Acerophagus papayae, the encyrtid
recommended. In case of severe incidence, parasitoid @ 100 per hamlet is recommended as
spraying of any one of the following chemicals the best management strategy (Jayaraj and
is recommended: Dimethoate (2 ml/litre), Ananthan 2008). After the complete control of
Profenophos (2 ml/litre), Chlorpyriphos (5 ml/ ants, release predatory ladybird beetle particu-
litre), Buprofezin (2 ml/litre), Imidacloprid (0.6 larly C. montrouzieri is to be considered to check
ml/litre), Thiamethoxam (0.6 g/litre). The insec- the mealybugs on cocoa in general.
ticide must be applied only after collecting the
pods which are ready for harvesting (Jayaraj and
Ananthan 2008). The possibility of exploiting the 58.4 Tea
use of natural enemies, semiochemicals, tolerant
or cocoa varieties unattractive to pseudococcids, The predator Cryptolaemus montrouzieri was
and the sterile male technique for effective vector released against the mealybugs on tea
control are to be considered (Padi 1997). (Dzhiviladze 1979). Cryptolaemus montrouzieri
was found colonizing on sucking insects in tea
gardens in China (Xuan Dai 1996). A new genus
58.3.6 Biological Control and species Assamencyrtus jorhatensis Singh
from Assam, India, is described. This species is
Indigenous natural enemies, though present in all recorded as a primary parasitoid of coconut-
situations, are not in sufcient numbers to lower mealy-bug, which has been observed as a serious
554 C. Mohan et al.
Table 58.3 List of mealybugs recorded on tea the tea gardens monitored, four generations per
Mealybug species Region/country References year are indicated by peaks in crawler density.
Crisicoccus India Williams (2004) Changes in the tea mealybug within tea tree dis-
matsumotoi tribution showed greater seasonal variation than
(Siraiwa) its density. The most notable aspect of the within-
Dysmicoccus India Williams (2004)
tree distribution is that some tea mealybugs were
impararlis sp.n.
found on the roots on all sample dates. The pro-
Lankacoccus Sri Lanka Williams (2004)
ornatus (Green) portion was smallest during the June-July period,
Nipaecoccus India Williams (2004) when the mealybug density was peaking (during
viridis (Newstead) Sri Lanka Williams (1999) the spring ush of growth and the harvest period).
Planococcus Ben-Dov (1994) As tea mealybug population entered the warmest
minor (Maskell) summer months the proportion of the population
Paraputo India Williams (2004) found under ground and on the lower portion of
theaecola (Green) the tea trunk increased. The tea mealybug was
Pseudococcus Sri Lanka Williams (2004) seeking protection (probably from heat) under-
theae (Rutherford)
ground (Abbasipour and Taghavi 2007).
Pseudococcus Iran Abbasipour and
viburni (Signoret) Taghavi (2007)
Rasrococcus India Ben-Dov (1994),
iceryoides (Green) Williams (2004) References
Rhizoecus theae Japan Ben-Dov (1994)
Kawai & Takai Abbasipour H, Taghavi A (2007) Description and sea-
sonal abundance of the tea mealybug, Pseudococcus
viburni (afnis) (Signoret) found on tea in Iran.
J Entomol 4(6):474478
Abraham EV, Padmanbhan MD (1967) Pests that damage
pest of young coconut nuts in Assam (Singh cocoa in Madras. Indian Hortic 11:1112
Abraham CC, Remamony KS (1979) Pests that damage
2006). Rhizoecus theae sp.n. was found infesting
cocoa plants in Kerala. Indian Cocoa Arecanut Spices
on the roots of tea in Japan (Kawai and Takagi J 2:7781
1971). In tea gardens of north of Iran, obscure Ackonor JB, Mordjifa DK (1999) Parasitism and preda-
mealybug Pseudococcus viburni (Ps. afnis) tion in Planococcoides njalensis (Laing) (Homoptera:
Pseudococcoidae) on Cocoa in Ghana. Trop Agric
(Signoret) was recorded as dominant species
76:269274
from tea gardens. Tea mealybug density increased Ali M (1987) Palm mealybug, Palmicultor palmarum
rapidly to an early peak in April, followed by a (Homoptera: Pseudococcidae), new to the Indian sub-
decline and then a low, but steady density for continent. Ann Entomol Soc Am 80(4):501
Anand B, Ayub Khan P (2006) Attractivenessof hibiscus
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cated by peaks in crawler density (Abbasipour First record of Eggplant Mealybug,
Coccidohystrixinsolita (Hemiptera: Pseudococcidae),
and Taghavi 2007). Nipaecoccus viridis
on Guam: potentially a major pest First record of Egg
(Newstead) was found mainly on the axils of plant Mealybug, Coccidohystrixinsolita (Hemiptera:
leaves and the growth of the affected shoots in Pseudococcidae), on Guam: potentially a major pest.
Assam (Das and Ganguli 1961). Two mealybugs, Data J 2:e1042 (23Jan2014)7/ BDJ.2.e1042 (le:///H:
/First%20record %20of%20 Eggpla.htm)
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insect pests of Bermuda. Bull Entomol Res (Coccidae), Part 3. With a catalogue of all species hith-
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Ferrero D (2008) Communication behaviors of the 6(12):3
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(Balachowsky 1957) (Pseudococcidae: Rhizoecinae) Theobroma cacao L. in Kerala with special reference
with its symbiotic ant Acropyga fuhrmanni (Forel to the mealy bug Planococcus lilacinus (Ckll.)
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pp 461465 horti_plantation%20crops_cocoa.html. Assessed
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5(3):245246 mealybugs (Hemiptera: Pseudococcidae). Bull
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virus-tolerant young cocoa. Report, Cocoa Research mealybug, Dysmicoccus cocotis (Maskell) and of a
Institute, Ghana, pp 252254 new related species on coconut in southern Asia
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pp 249263 Rutherford from Sri Lanka and Trionymus kayashimai
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(2003) New report of mealybugs occurring inside the Entomol Month Mag 135(16161619):9193
perianth of immature nuts in coconut. Insect Environ Williams DJ (2004) Mealybugs of southern Asia. The
9(2):5354 Natural History Museum/Southdene SDN. BHD,
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pests of arecanut and their control. Arecanut J 12:136 Williams DJ, Granara de Willink MC (1992) Mealybugs
Razak TA, Jayaraj S (2002) Incidence of spiralling white- of Central and South America. CAB International,
y and leaf mealybug on coconut. Insect Environment London. 635p
8(1):3839 Williams DJ, Matile-Ferrero D (2005) Mealybugs from
Roivainen O (1976) Transmission of cocoa viruses by Zanzibar and Pemba islands with a discussion of a
mealybugs (Homoptera:Pseudococcidae). J Sci Agric potential invasive species (Hemiptera,
Soc Finl 48(3):203304 Pseudococcidae). Rev Fr dEntomol 27(4):145152
Singh S (2006) A new encyrtid genus Assamencyrtus Xuan Dai (1996) Investigation on ladybird beetles
(Hymenoptera: Chalcidoidea), from Assam, India, (Coccinellidae) in tea gardens of east Guizhou. J Tea
parasitising coconut mealybugs (Homoptera: Sci 16:131134
Rubber
59
Mani Chellappan
Mealybugs are injurious to rubber (Hevea brasil- Both crawlers and adult female mealybugs feed
iensis) in Sri Lanka, India, Indonesia, Egypt, on the sap of rubber plants by inserting their sty-
Malaysia, Nigeria, Tanzania, etc. (Table 59.1). lets into the epidermis of the leaf, tender shoots,
Mealybug infestation had not been a major prob- main stem, inorescence, as well as the fruit. Due
lem on rubber until the recent introduction of the to the feeding, the leaves showed chlorosis, plant
mealybug Paracoccus marginatus in India. stunting, leaf deformation, early leaf and fruit
Among the mealybugs infesting rubber, except P. drop, heavy buildup of honeydew and consequent
marginatus, all are considered to be minor. sooty mould development on all plant parts, and
Ferrisia virgata were found attached to the roots drying of worst affected branches (Mani
hanging down from an African rubber tree (http:// Chellappan 2010).
bugguide.net/node/view/148641/bgpage).
M. Chellappan (*)
Kerala Agricultural University,
Thrissur 680656, Kerala
e-mail: mani-chellappan@kau.in
and around the rubber plantation, viz., %) or neem oil emulsion (12 %) as spray were
Eupatorium odoratum (Chromolena oderata), recommended to control the mealybugs.
Berrari sp., Lantana aculeata, Mimosa pudica, Application of thiamethoxam 25 %WG (@ 1 g/L,
lmpertala cylindrica, and a variety of other plants imidacloprid 480 SL (@ 1 mL/L), acetamiprid 20
had papaya mealybug infestation, including glyr- % (1 g in 1 L of water), and mineral oil (@ 5 mL/L)
icidia. Mulching with dry leaves, grass cuttings, was recommended to control P. marginatus
and cover crop looping around the plant, is rec- (Galanihe et al. 2010). Acephate, carbaryl, chlor-
ommended as a cultural operation for rubber pyrifos, diazinon, dimethoate, malathion, and
nurseries, and young seedlings also aggravated white mineral oils (Mani Chellappan 2010),
the problem. Also, the abnormal leaf fall in rub- dimethoate EC 40 % (1 mL in 1 L of water) / imi-
ber caused by the fungi Phytophthora palnivora dacloprid 20 % (1 mL in 1 L of water)/thiameth-
during wet weather coupled with humid condi- oxam 25 % (12 g in 1 L of water)/ acetamprid 20
tion and powdery mildew disease caused by fun- % (1 g in 1 L of water) (http:// www.plantationin-
gus Oidium heveae steinm on newly formed dustries.gov.lk/ dwnlds/ plantation/8.pdf) were
tender ush during the refoliation period of recommended to control P.marginatus on rubber.
January to March (symptoms including tender
leaves with ashy coating curl, crinkle, edges roll
inward and fall, leaving the petit. Die-back of 59.7 Biological Control
twigs also follows. On older leaves, white patches
appear causing necrotic spots affecting ower Biological control includes spraying entomo-
and tender fruits, which are shed affecting seed pathogenic fungus, viz., Verticilium leccanii (10
production) suppressed the actual symptom of g/L) in 5 % soap solution, encouraging the lepi-
the mealybug on rubber (Mani Chellappan 2010). dopteran predator, Spalgis epeus and release of
aphelinid Acerophagus papayae (Noyes and
Schauff). The parasitoid was found highly suc-
59.5 Management cessful in suppressing P. marginatus on rubber.
Kerala accounts for 92 % of the total area in India
Cultural management includes inspecting all under rubber cultivation, that is, 5.17lakhs ha.
papaya, temple trees, and other susceptible hosts Average production is 1949 kg/ha. Average price
in and around rubber plantations; burning and is Rs. 1619/kg; Yield reduction 10 %; Total
destroying the severely infected trees/parts of infested area49,500 ha; Saving Rs. 17.85 lakhs
trees immediately; avoiding transportation of (Mani Chellappan 2010).
infested plant material; avoiding pruned, infested
plant parts being left unattended or being placed
in garbage bins or vehicles; washing the insects References
off the plants with a powerful water jet; wrapping
polythene/spongy tapes impregnated with insec- Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
ticides around tree trunks to exclude ants from
Pseudococcidae and Putoidae) with data on geograph-
the canopy; and unsettling the crawlers with a jet ical distribution, host plants, biology and economic
of water. importance. Intercept Limited, Andover, 686p
Chellappan M (2010) Status of papaya mealybug,
Paracoccus marginatus Williams and Granara de
Willink in Kerala. In: Shylesha AN, Joshi S, Rabindra
59.6 Chemical RJ, Bhumannavar BS (eds) Classical biological con-
trol of papaya mealybug, Paracoccus marginatus in
Spraying soap solution (5 %) to dissolve the wax India Proceedings of the national consultation meet-
ing on strategies for deployment and impact of the
and expose the mealybug body to various methods
imported parasitoids of papaya mealybug. NBAII
of management and use of tobacco decoction (2 (ICAR), Begaluru, pp 4042
560 M. Chellappan
Cox JM (1989) The mealybug genus Planococcus Jayasinghe CK (1999) Pests and diseases of hevea rubber
(Homoptera: Pseudococcidae). Bull Br Mus Nat Hist and their geographical distribution. Bull Rubber Res
Entomol 58(1):178 Inst Sri Lanka 40:18
Galanihe LD, Jayasundera MUP, Vithana A, Lyla KR, Philip BM (2010) Incidence of papaya mealy-
Asselaarachchi N, Watson GW (2010) Occurrence, bug, Paracoccus marginatus Williams and Granara de
distribution and control of papaya mealybug, Willink in Kerala. Insect Env 15(4):156
Paracoccus marginatus (Hemiptera: Pseudococcidae), Mathew J (2011) Status of papaya mealy bug on rubber in
an invasive alien pest in Sri Lanka. Trop Agric Res Ext Kerala. In: Proceedings of the National consulation
13(3):8186 meeting on strategies for deployment and impact of
Hall WJ (1921) The hibiscus mealybug. Ministry Agric the imported parasitoids of papaya mealybug,
Egypt Tech Sci Ser Entomo Sec Bull 17:128 Classical biological control of papaya mealybug
Hill DS (2008) Pests of crops in warmer climate and their (Paracoccus marginatus) in India, p 60
control. Springer, Dordrecht, 625p Venette RC, Davis EE (2004) Mini risk assessment pas-
Iheagwam EU (1981) Natural enemies and alternative sionvine mealybug: Planococcus minor (Maskell)
hostplant of the cassava mealybug, Phenacoccus [Pseudococcidae: Hemiptera]. Department of
manihoti (Homoptera, Pseudococcidae) in south- Entomology, University of Minnesota, St. Paul, 30p
eastern Nigeria. Revue de Zool Afr 95(2):433438 Williams DJ (2004) Mealybugs of southern Asia. The
Natural History Museum/Southdene SDN. BHD,
London/Kuala Lumpur, 896 p
Cashew
60
V. Ambethgar
Mealybugs are injurious to cashew plantations in (Ambethgar 2011). In South India, the rst out-
India, West Africa, Tanzania, Thailand, etc. break of F. virgata on cashew was discovered
(Table 60.1). during FebruaryMarch 1998 in Cuddalore dis-
In India, severe incidence of mealybugs was trict of Tamil Nadu. Ferrisia virgata is largely
observed on cashew in Maharashtra (Godse et al. dominant across the cashew-growing areas
2003), Tamil Nadu (Ambethgar et al. 2000), (Ambethgar et al. 2000).
Kerala, Karnataka, and Andhra Pradesh
V. Ambethgar (*)
Tamil Nadu Agricultural University,
Regional Research Station,
Vridhachalam 606 001, Tamil Nadu, India
e-mail: drva1965@gmail.com
The plants phloem, which contains the nutri- of black sooty mold fungi (Capnodium species)
ents needed for mealybug development. As that can result in further plant damage, because it
mealybugs digest their food, they excrete sugar- hastens the germination of sooty molds, which
rich uid called honeydew. Mealybugs also block light from the leaves and impede photosyn-
exude brous, white waxy beard that hangs from thetic efciency of plants. The honeydew and
the tree trunk and branches. In cashew, mealybug sooty mold contamination may also impair the
infestations are more often conned to short peri- quality of the cashewnut (Ambethgar et al. 2000).
ods, synchronizing with ushing through the Additionally, the feeding punctures resulting
fruiting season (February-May) rather than the from mealybug infestation facilitate secondary
remaining periods of the dormant season. Both infection of twig-blight disease caused by
nymphs and adults prefer to feed on the tender Pestalotia microspora (Ambethgar 2011).
shoots, nodes, petioles, leaves, inorescence, Severe outbreaks of mealybug have been
ower panicles, or developing fruit clusters reported on cashew, adversely affecting the yield
which are soft and succulent. Mealybugs, while of cashewnut over 80 % in extreme cases in the
desapping the plants, inject salivary toxins, which state of Tamil Nadu, India. Aapparent yield
resulted in malformed leaves, reduced plant losses in terms of raw cashewnut have been
vigor, stunted growth, and occasional death of reported to be varying from 7 to 16 % and 2350
branches. Severely infested cashew trees could % during 19981999 and 20032004, respec-
be easily sighted even from a distance by their tively (Ambethgar 2011). Such a massive yield
sickly appearance. On susceptible trees, affected losses occur when mealybugs infest fruit clusters
leaves showed a characteristic curling/premature or excrete honeydew that covers fruit and
senescence (yellowing), similar to the damage foliage.
caused by viruses. If ower blossoms were
attacked, the fruits could set poorly. When fruits
are infested, they could be entirely covered with 60.2 Seasonal Occurrence
the waxy white coating of the mealybug. Heavy of Mealybugs
infestation could lead to fruit rot drop, or fruits
remained on the bunch in a shriveled and dry Temperature is the driving force for mealybug
condition. Sometimes, the whole plants may wilt upsurge. The maximum mealybug population
and become stunted. New growth may become was found when dry and humid conditions pre-
distorted. vail from March to May. There has been positive
More importantly, the direct sap feeding on correlation between temperature and mealybug
developing fruit clusters results in ill-lled nut population, and also signicant negative correla-
meal/kernel formation, improper shell-split, and tion with relative humidity and total number of
reduced shelling outturn, which ultimately impair rainy days on mealybugs. The population of F.
the market value of cashewnuts. Besides, mealy- virgata is abundant during JanuaryMay, coin-
bug causes indirect physical damage as they ciding with active new ushing, owering, and
excrete the carbohydrate-rich clear sticky honey- fruit development periods of cashew in cashew
dew which can accumulate on the foliage and orchards. Four to ve generations are completed.
fruit clusters, and supports the growth of black The rapid population increase in summer is fol-
sooty mold fungus. Though honeydew can be lowed by an equally rapid decline after harvest.
dissolved even by a slight rain, they readily dry in Low precipitation of Northeast seasonal mon-
warm temperatures. When mealybug populations soon (OctoberDecember) is found to be favor-
are severe, honeydew can accumulate to form a able for the early buildup of mealybugs under the
hard, wax-like layer that covers the infested climatic conditions of Tamil Nadu. Once
plant, which clog stomatal openings and impede Southwest monsoon (MayJune) sets in due to
gas exchange and respiration. Undoubtedly, hon- heavy downpour of rain, mealybugs are washed
eydew serves as a substrate for the development out. Due to increase in humidity and decrease in
564 V. Ambethgar
the temperature, the mealybug population Diacamma rugosum (Le Guill), Monomorium
declines and becomes almost negligible. latinode (Forel), Oecophylla smaragdina
Continuous monsoon, low temperature, and (Fabricius), and Technomyrmex sp. (Ambethgar
humidity are detrimental for mealybug develop- 2002b) were found to be associated with mealy-
ment (Abdul Rahiman et al. 1995). bugs in cashew ecosystem. Mealybugs are known
to bribe ants with their sugary secretion called
honeydew, and in return, ants help in the spread-
60.3 Association of Ants ing of mealybugs and provide them protection
with Mealybugs from predators, parasitoids, and other natural
enemies (Bentley 1977). Most often, colonies of
Four ant species, viz., Anoplolepis longipes M. hirsutus are attended by red weaver ant
(Jerdon), A. gracilipes (Smith), Tapinoma indi- Oecophylla smaragdina. These ants are known to
cum (Ingar), T. melanocephalum (Fabricius) attack the natural enemies of mealybugs while
(Rickson and Rickson 1998), and seven species, attending the pests. Thus, ants can exacerbate
namely Camponotus compressus (Fabricius), C. mealybug pest problems by disrupting the natural
sericeus (Fabricius), Crematogaster sp., enemy activity in cashew.
Prevention is better than cure. This principle is Farm-level regular monitoring, early detection,
highly applicable in the management of cashew and isolation of infested plants are important to
mealybugs. Mealybugs are best treated if detected avoid outbreaks. Visual plant inspection is an
early, when populations are low. Once they efcient way to detect early mealybug infesta-
become established, mealybugs are very difcult tions. Early infestations can be easily overlooked
to achieve effective control. Hence, a concomi- due to the mealybugs tendency to hide in pro-
tant use of pest monitoring, cultural, mechanical, tected locations. Presence of white ecks or waxy
biological, and chemical methods of control at residues along the leaf midribs, on leaf or stem
appropriate time have to be integrated for long- axils, and on the underside of leaves is an indica-
term management of mealybugs and to reduce tion of mealybug infestation. Honeydew, sooty
the yield loss. mold, and the presence of ants may also be indi-
cations of mealybug infestation.
60 Cashew 565
virgata on cashew (Ambethgar et al. 2000). Nadu. In: Mason PG, Gillespie DR, Vincent C (eds)
Proceedings of ISBCA 3, USDA-Forest Service,
Sprays of malathion (0.05 %) or monocrotophos
pp 562563
(0.05 %) or phosphamidon (0.03 %) or dimetho- Ambethgar V (2011) Field evaluation of some insecti-
ate (0.03 %) were recommended for the control cides against white-tailed mealybug, Ferrisia virgata
of F. virgata on cashew (Ambethgar 2011). Use (Cockerell) infesting cashew. In: Souvenir and abstract
of the international symposium on Cashew, 0912
of chlorpyriphos 20 EC @ 2.5 mL/L offered ade-
December 2011, Madurai, India, pp 131132
quate control of mealybugs in cashew. Foliar Ben-Dov Y (1994) A systematic catalogue of the
sprays of dichlorvas (0.05 %) was most effective, mealybugs of the world (Insecta: Homoptera:
recording 93.5 % mealy bug reduction over con- Coccoidea: Pseudococcidae and Putoidae) with data
on geographical distribution, host plants, biology
trol in 10 days after rst spray, and contributed to
and economic importance. Intercept Limited,
a maximum nut yield (1200 kg/ha (Ambethgar Andover, 686 p
2011). Triazophos (0.05 %) and profenophos Bentley BL (1977) The protective function of ants visiting
(0.05 %) were at par and recorded 91.292.5 % the extraoral nectaries of Bixa orellana L. (Bixaceae).
J Ecol 65:2738
reduction of mealybug over control, and contrib-
Chellappan M, Lawrence L, Indhu P, Cherian T, Anitha S,
uted to the nut yield of 11001125 kg/ha. Jimcymaria T (2013) Host range and distribution pat-
Currently, newer insecticides in the group of tern of papaya mealy bug, Paracoccus marginatus
neonicotinoids with more novel modes of action Williams and Granara de Willink (Hemiptera:
Pseudococcidae) on selected Euphorbiaceae hosts in
have also gained in popularity for control of
Kerala. J Trop Agric 51(12):5159
mealybugs. Thiamethoxam (0.003 %) and imida- De Lotto G (1964) Observations on African mealy bugs
cloprid (0.005 %) were at par and recorded 91.2 (Hemiptera: Coccoidea). Bull Brit Mus Nat Hist
92.5 % reduction of mealybug over control with Entomol 14:343397
Godse SK, Bhole SR, Munj AY, Gurav SS (2003)
nut yield of 11001125 kg/ha (Ambethgar 2011).
Chemical control of mealybugs (Ferrisia virgata
Insecticides recommended for control of mealy- Cockerell). Cashew 17(2):1517
bugs in cashew production are carbaryl 50 WP Lad SK, Patil PD, Godase SK (2013) Record of mealy
(2.0 g/L), acephate 75 SP (1.0 g/L), chlorpyri- bugs infesting fruit crops in in Konkan Region of
Maharashtra. J Appl Zool Res 24(2):141145
phos 20 EC (2.5 mL/L), dichlorvas 76 WSC (1.5
Mahapatro GK (2008) Helopeltis management by chemi-
mL/L), dimethoate 30 EC (2.0 mL/L), malathion cals in cashew: A critical concern. Indian J Entomol
50 EC (2.0 mL/L), monocrotophos 36 WSC (1.5 70(4):293308
mL/L), phosphamidon 75 EC (1.0 mL/L), pro- Mani M, Krishnamoorthy A, Singh SP (1990) The impact
of the predator, Cryptolaemus montrouzieri Mulsant,
fenophos 40 EC (1.5 mL/L), triazophos 40 EC
on pesticide-resistant populations of the striped mealy-
(2.0 mL/L), imidacloprid 17.8 SL (0.5 mL/L), bug, (Ckll.) on guava in India. Insect Sci Appl
thiamethoxam 25 WG (0.5 mL/L), and azadi- 11(2):167170
rachtin 0.03 EC (3.0 mL/L). Maniania NK (2011) Integrated management of major
insect pests and diseases of cashew in East and
Western Africa. Factsheet-BAF Advisory Service on
Agricultural Research for Development, Gotingen,
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Maruthadurai R, Desai AR, Chidananda Prabhu HR,
Abdul Rahiman P, Vijayalakshmi CK, Reddy AGS (1995) Singh NP (2012) Insect pests of cashew and their man-
Occurrence and distribution of mealy bug in coffee. agement. Technical Bulletin 28/2012, ICAR Research
Kisan World 22(11):3940 Complex for Goa, India, 16 p
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(2000) Managing mealybugs in cashew. Science and Publication, New Delhi, 124p
Technology, The Hindu, February 24, 2000 Rickson FR, Rickson MM (1998) The cashew nut,
Ambethgar V (2002a) Record of entomopathogenic fungi Anacardium occidentale (Anacardiaceae), and its
from Tamil Nadu and Pondicherry. J Entomol Res perennial association with ants: extraoral nectary
26(2):17 location and the potential for ant defense. Am J Bot
Ambethgar V (2002b) Insect visitors of cashew in North- 85(6):835849
Eastern Zone of Tamil Nadu. Progress Hortic Strickland AH (1947) Coccids attacking cacao
34(2):223229 (Fheobroma cacao L.), in West Africa, with descrip-
Ambethgar V, Bhat PS (2008) Entomogenous fungi asso- tions ofve new species. Bull Entomol Res
ciated with insect pests in cashew orchards of Tamil 38:497523
60 Cashew 567
Sunitha ND, Jagginavar SB, Biradar AP (2009) Entomology, University of Minnesota, St.Paul, USA,
Bioefcacy botanicals and newer insecticides against pp 130
grape vine mealybug, Maconellicoccus hirsutus Williams DJ (1996) A synoptic account of the mealybug
(Green). Karnataka J Agric Sci 22:710711 genus Ferrisia (Hem., Pseudococcidae). Entomol
Topper CP (2002) Issues and constraints related to the Month Mag 132:110
development of cashew nuts from ve selected African Williams DJ (2004) Mealybugs of southern Asia. The
countries. International Trade Centre (ITC)-Common Natural History Museum/Southdene SDN. BHD,
Fund for Commodities (CFC), Reunion Regionale sur London/Kuala Lumpur, 896 p
le Developpement des Exportations de noix de Cajou Williams DJ, Matile-Ferrero D (2005) Mealybugs from
d Afrique, Cotonou, Bnin, pp 124 Zanzibar and Pemba islands with a discussion of a
Venette RC, Davis EE (2004) Mini risk assessment: pas- potential invasive species (Hemiptera,
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Oil Palm
61
P. Kalidas
P. Kalidas (*)
Directorate of Oil Palm Research,
Pedavegi 534450, Andhra Pradesh, India
e-mail: pothinenikali@redif.com
Mealybugs are injurious to spice crops like black observed in all the taluks surveyed in Wayanad
pepper, cardamom, ginger, turmeric, etc. The (Kerala). The pest infestation was also observed
mealybug species recorded on various spice in Udumbanchola (Idukki district), Kozhikode
crops in different regions are listed in Table 62.1. (Kozhikode district), and Taliparamba (Kannur
district) taluks in Kerala. The pest infestation was
higher in Wayanad (8.021.1 %) and lower in
62.1 Black Pepper (Piper nigrum) Idukki (03 %). Stray infestations of the pest
were also observed in Kozhikode and Kannur dis-
In Indonesia, Planococcus citri and Ferrisia vir- tricts in Kerala. Among the taluks in Kerala, the
gata are reported to infest the aerial parts (shoots, percentage of vines infested by root mealybugs
leaves, spikes, and berries) of black pepper vines was higher in Vythiri (21.1 %) taluk. In Karnataka
(Kueh et al. 1993). In Sri Lanka, P. citri is reported state, mealybug infestation was conned to
to infest the roots of black pepper vines Kodagu and Hasan districts (Alur and Saklespur).
(Dharmadasa 2000), whereas in China, Mealybug infestation was higher in Kodagu (1.7
Planococcus sp. and P. lilacinus in Vietnam 15.1 %) district and lower in Hassan (04.4 %)
are known to infest the roots of black pepper vines district. Among the taluks in Kodugu district, the
(Sarma 2010). Twelve species of mealybugs are percentage of vines infested by root mealybugs
known to infest black pepper in India, and among was higher in Virajpet (15.1 %) taluk. Based on
them, the root mealybugs are the most severe the distribution pattern, it was concluded that a
(Devasahayam et al. 2009). According to these highly signicant and positive correlation
authors, the mealybugs Planococcus sp., (r = 0.451**) was observed between the pest
Planococcus citri, Planococcus lilacinus, Ferrisia infestation and altitude of the location. A mean of
virgata, and Dysmicoccus brevipes are known to 0.1 % of vines were infested at lower altitudes
infest the roots and basal regions of black pepper (0250 m above MSL) when compared to 8.9 %
vines in India and were conned to certain parts at higher altitudes (7511000 m above MSL).
of Kerala and Karnataka states. In Kerala, root
mealybug infestations on black pepper were
62.1.1 Damage
S. Devasahayam (*) T.K. Jacob In India, the incidence of mealybugs infesting the
Indian Institute of Spices Research, aerial parts of the vine (all species combined)
Marikunnu P. O., Kozhikode 673 012, Kerala, India was highest in Kasargod district (Kerala),
e-mail: devasahayam@spices.res.in
followed by Idukki district (Kerala), wherein 2.7 were covered with mulch. The pest infestation
and 2.3 % of leaves of affected vines (5.4 %) results in defoliation, yellowing and wilting of
were infested (Koya et al. 1996). The aerial infes- leaves and lateral branches, and also mortality of
tation of mealybugs such as F. virgata and P. citri vines in severe cases of infestation (Devasahayam
is mainly seen on tender shoots, leaves and et al. 2009). Various intercrops (coconut,
spikes, especially in the nursery. Planococcus arecanut, coffee, banana, colocasia, cardamom,
minor, P. longispinus, and P. orchidicola are gen- and turmeric) grown in black pepper gardens
erally encountered within old leaf galls induced were found infested with root mealybugs. Root
by leaf gall thrips (Liothrips karnyi Bagn.), prob- mealybug infestations were observed on black
ably due to the conducive microclimatic condi- pepper vines trailed on all standards (support
tions within them (Devasahayam 2000). In trees like silver oak, Erythrina spp., and jack-
nursery plants, infestations by F. virgata and fruit) (Devasahayam et al. 2009). Continuous
P. citri result in wilting of the affected parts. infestation without proper management leads to
gradual decline and death of the vine. The infes-
tation is generally greater during the post-mon-
soon season and lesser during summer months.
62.1.2.1 Pathogens
The fungus Phytophthora capsici (Leonian) and
nematodes such as Meloidogyne incognita
(Kofoid and White) (Chitwood), and Radopholus
similis (Cobb) were commonly associated with
Ferissia virgata on black pepper root mealybug infested vines. At Wayanad and
Kozhikode districts, all the root mealybug
Colonies of root mealybugs are distributed on infested vines examined (n = 104) were also
the main, secondary, and tertiary roots, basal infested with either Phytophthora and nematodes
region of stems on rooted cuttings in the nursery, or both. The infested vines exhibited symptoms
and also on the vines of all age groups in the eld. such as rotting of roots, absence of feeder roots,
The mealybug colonies are observed even up to a yellowing and wilting of leaves, defoliation, and
depth of 2 ft below the soil in severely affected mortality of vines that are characteristically asso-
vines. The infestation on the basal regions of the ciated with P. capsici and nematode infections.
stem is seen under the soil and also when they At a few locations in Wayanad district, the root
576 S. Devasahayam and T.K. Jacob
The fungus Diacanthodes philippinensis tricts in Kerala and Karnataka (Bhat et al. 2003a,
(Pat.) (Singer) is reported to be associated with P. b, 2005)
lilacinus on coffee in India, and whenever both
occurred together, the plants wilted and died.
Infestation by the mealybug alone did not cause 62.1.3 Ants
the death of coffee plants (Sekhar 1964; Chacko
and Sreedharan 1981). In Africa, the coffee root Three species of ants, namely, Anaplolepis sp.,
mealybug, which was earlier identied as P. citri Crematogaster sp., and Technomyrmex sp., and
and associated with the fungus D. novoguineesis two unidentied species were associated with
(Hennings) Fidalgo, has been later described as a root mealybug colonies. In many cases, it was
new species, P. fungicola (Watson and Cox easier to identify infested vines based on the
1990). The fungus is considered as a symbiont activity of the ants (Devasahayam et al. 2009).
providing protected cavities on the root surface in
which the mealybugs live in return for the sugars
in the honeydew excreted by them and in the sap 62.1.4 Natural Enemies
that escapes from the insects feeding punctures in
the roots (Fidalgo 1962). On black pepper, Leptacis sp. (Platygasteridae)
Ferrisia virgata and P. citri were identied to and Blepyrus insularis (Cam.) (Encyrtidae) were
transmit Piper Yellow Mottle Virus (PYMoV), found parasitizing Pseudococcus sp. and F. vir-
the badnavirus, causing stunt disease which is gata, respectively, infesting on the aerial parts of
increasingly becoming serious, especially at the vine (Devasahayam and Koya 1998). Larvae
higher altitudes in Wayanad and Kodagu dis- of Spalgis sp. (Spaligidae) were observed to
62 Spices 577
predate on root mealybug colonies, especially three times a season, has been recommended.
those at the base of the stems (Devasahayam Use of Erythirna variegata (L.) and E. orientalis
et al. 2009). (Murr.) as live supports is not advocated (Sarma
2010). In Indonesia, spraying albolineum (200
250 mL in 18 L water) or dimethoate (3540 mL
62.1.5 Management in 18 L water) or malathion (45 mL in 18 L water)
has been suggested for controlling P. citri and F.
virgata (Kueh et al. 1993).
Microbial Pathogens The fungal pathogen
Metarhizium anisopliae (Metsch.) (Sorokin) was 62.1.5.1 Integrated Management
found to cause 79.6 % reduction in mealybug In India, an integrated pest management strategy
population, 30 days after treatment under labora- for the management of root mealybugs was
tory conditions (Devasahayam and Koya 2000). developed based on eld trials conducted with
promising insecticides and plant products. The
Natural Products Alcoholic extracts (3 %) of strategy involves planting root mealybug-free
Azadirachta indica and Vitex negundo; tobacco rooted cuttings in the eld, removal of weeds in
extract (3 %); custard apple seed extract (2 %); the interspaces of black pepper vines during sum-
and agro spray oil (3 %) are known to cause up to mer, drenching tobacco extract (3 %) or custard
75 % reduction in root mealybug population, 30 apple seed extract (2 %) on mildly affected vines,
days after treatment. Among the neem products, or drenching imidacloprid (0.0125 %) or aceta-
Nimbicidine (0.5 %) was the most effective, maprid (0.0125 %) or carbosulfan (0.075 %) or
resulting in 60 % reduction in the population of chlorpyriphos (0.075 %) on the affected vines,
root mealybugs, 30 days after treatment and adoption of control measures against
(Devasahayam 2006). Phytophthora and nematode infections
(Devasahayam 2006). Adequate care should be
Insecticides In India, imidacloprid (0.0125 %), taken to ensure that the insecticide solution per-
acetamaprid (0.0125 %), and carbosulfan colates down to the roots while drenching the
(0.075 %) were more promising, resulting in over vines.
90 % reduction in the population of root mealy-
bugs, 30 days after treatment under laboratory
conditions (Devasahayam 2006). Dimethoate, References
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base of the plant with pronil 50 G-SC (5 mL in cotton agroecosystem of Punjab, Pakistan. Int J Agric
Biol 11:163167
10 L of water) and applying carbofuran 3G (10
Ben-Dov Y (1994) A systematic catalogue of the mealy-
15 g per vine) (Dharmadasa 2000), or drenching bugs of the world (Insecta: Homoptera: Coccoidea:
the base of the vine with chlorpyriphos (28 mL in Pseudococcidae and Putoidae) with data on geograph-
12 L water) (Sarma 2010) have been recom- ical distribution, host plants, biology and economic
importance. Intercept Limited, Andover, 686 p
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Entomon 34:147154
Mulberry
63
J.B. Narendra Kumar, M.A. Shekhar,
and Vinod Kumar
Mealybugs are injurious to mulberry (Morus Maconellicoccus hirsutus was rst detected on
spp.) in several countries (Table 63.1). Mulberry mainland US in August, 1999 in Imperial Valley,
is the sole food plant of the silkworm, Bombyx a low desert region in southern California on
mori L., the producer of fabulous silk which mulberry (Roltsch et al. 2006).
derives all the nutrients for its growth from the
mulberry leaf. Mealybugs pose serious threat to Seasonal Development In south India, though
mulberry cultivation mainly in India. Mulberry the pest is observed to infest mulberry throughout
fruits are edible, and can be great for health in the year, the incidence was highest during sum-
some countries. Production of appreciable quan- mer (34.93 %) and least during winter (9.45 %)
tity of quality mulberry leaf is hampered by the with an average incidence of 22.15 % (Hemalatha
mealybugs in silk-producing states in India. and Shree 2008; Sathya Prasad and Manjunath
Among the mealybug species, Maconellicoccus 1992). According to Narendra Kumar et al.
hirsutus and Paracoccus marginatus in plains (2006), in Bangalore rural district, the mealybug
and Paraputo sp. in the hilly regions caused dras- incidence reached its peak during April (18.79
tic reduction in mulberry leaf yield thereby %) and gradually started declining afterwards,
affecting the silk industry. and lowest incidence of 2.56 % was recorded
during December. Pink mealybug passes through
1015 generations in a year and was found active
63.1 Pink Hibiscus mealybug, even during winter months without any hiberna-
Maconellicoccus hirsutus tion (Rajadurai 2005b). Further, it completes its
life cycle in 2429 days on mulberry (Misra
In India, Misra (1919) reported for the rst time, 1919). However, Dhahira Beevi (1989) reported
the attack on mulberry by M. hirsutus. Though that total life span of the mealybug was 30.6 days
the pest is known to attack mulberry almost since for female while it was 22.7 days for males on
a century, it has assumed the key pest status, mulberry.
especially in Karnataka, Tamil Nadu and Andhra
Pradesh only about one and a half decade back Damage Mealybugs cause damage to mulberry
(Sriharan et al. 1979; Baskaran et al. 1994). crop by sucking the sap from young leaves and
buds. As mealybugs suck and feed, they inject
J.B.N. Kumar (*) M.A. Shekhar V. Kumar into the plant a toxic saliva that results in mal-
Central Sericultural Research and Training Institute, formed leaf and shoot growth, stunting and occa-
Mysore 570 008, India sional death (Lavanya Latha et al. 2004; Rajadurai
e-mail: jbnarendra@gmail.com
Table 63.1 List of mealybug species recorded on mulberry in different regions in the world
Species Region/country References
Atracoccus fuscus (Borchsenius) Turkmenistan Ben-Dov (1994)
Crisicoccus maricola Tang Mongolia Ben-Dov (1994)
Ferrisia virgata (Ckll.) Egypt Attia (2006)
Formicoccus lateens sp.n India Williams (2004)
Maconellicoccus hirsutus Green India Misra (1919), Raichoudhury (1958),
Manjunath et al. (1996)
China Snchez (2000)
Brazil de Almeida and Fonseca (2000)
Egypt Hall (1926)
Canada Garland (1998)
California Roltsch et al. (2006)
Philippines Mundo (1984)
Pakistan Zaman et al. (1996); Sahito et al.
(2012)
Bangladesh Ali and Ahmed (1990)
Iran Fallahzadeh et al. (2002)
Nipaecoccus vastator (Maskell) Iraq El-Haidari et al. (1978)
Niapecoccus viridis (Newstead) Pakistan Williams (2004)
Paracoccus marginatus Williams and India Mani Chellappan et al. (2013);
Granara de Willink Mahalingam et al. (2010); Shekhar
et al. (2011); Prasad et al. (2012)
Paraputo sp. India Misra et al. (1996); Biswas et al.
(2002)
Peliococcus mesaiaticus Borchsenius Afghanistan Ben-Dov (1994)
& Kozarzhevskaya
Phenacoccus divericatus sp.n. Pakistan & India Williams (2004)
Planococcus citri (Risso) Pakistan Williams (2004)
Planococcus minor Maskell Ben-Dov (1994)
Pseudococcus comstocki (Kuw.) California Bartlett and Clancy (1972)
Central Asia Kryachko (1978)
Armenia Oganesyan and Babayan (1979)
Crimea (USSR) Romanchenko and Belskaya (1981)
Georgia Kanchaveli and Partsvaniya (2009)
Pseudococcus longispinus (Targioni India Sakthivel et al. (2011)
Tozzetti)
Pseudococcus maritimus (Ehrhorn) Nearctic & neotropical Ben-Dov (1994)
Spilococcus mari (Siraiwa) Japan Ben-Dov (1994)
Trionymus mori Lobdell Mississippi Ben-Dov (1994)
2005a). The damage by the pink mealybug gives mity symptom caused due to pink mealybug
disease like appearance called as tukra. A heavy infestation.
black sooty mould develops on the infested leaves
and stem as a result of honey dew excretions by The extent of damage by the pink mealybug in
the mealybug. Earlier tukra in mulberry was mis- mulberry is reported to be 34.24 % (Manjunath
taken for a viral disease and mealybugs were et al. 1996) leading to an estimated leaf yield loss
believed to be the vectors of the same of about 4,500 kg/ha/year with a cocoon crop loss
(Rangaswamy et al. 1976). Later it was discov- of about 1015 % (Manjunath et al. 2000;
ered that so-called tukra disease is only a defor- Rajadurai and Thyagarajan 2003). Due to this, the
63 Mulberry 581
sericulturists are constrained to forego a rearing of and least in Dharmapuri (12.06 %) and Thanjavur
about 450 layings/ha/year, thus reducing cocoon districts of Tamil Nadu, India (Baskaran et al.
production by about 300350 kg/ha/year (Kumar 1994). Mealybug infestation had resulted in 30 to
et al. 1995) which works out to be Rs. 60,000 40 % loss in mulberry leaf yield (Nighat Mehmood
70,000 annually. Severe incidence of the mealy- 2004). In West Bengal, a leaf yield loss of 7.95 to
bug has been reported in Erode district (63.12 %) 11.03 % has been reported (Anonymous 2011).
extract and Pongamia seed kernel extract were single dose and irrigated once in 6 days than
found to be more effective than seed oils against applying nitrogenous fertilizer as a split dose and
mealybug (Narendra Kumar et al. 2006). providing irrigation either once in 6 days or 8
Spraying with 0.03 % Azdirachtin was recom- days.
mended for the mealybug control @ 5 ml/litre
(safety period: 10 days) (Anonymous 2010a). Mechanical Method The mechanical control of
mealybugs includes clipping of infested portion
Cultural Method Kasi Reddy et al. (2004) by sickle or secateur, collecting them in a poly-
reported that raising of maize as intercrop in mul- thene bag or bucket and destroying them by burn-
berry plantation increased the population of pred- ing or dipping in 0.5 % soap solution (Rajadurai
ator, Cheilomenes sexmaculata, (Fabricius) 2005a). According to Tomy Philip et al. (2002),
doubly (44 %) compared to the mulberry without chopping the affected portion and killing the
intercrop (21 %) resulting in the suppression of mealybugs either by burning or dipping them in
the pink mealybug population by 84 %. Growing 0.5 % DDVP with 0.5 % soap solution after prun-
cowpea as intercrop with mulberry enhances the ing or leaf harvest was found to be effective in
population of predatory ladybird beetles such as reducing mealybug population in mulberry.
C. sexmaculata, which initially feeds on cowpea
aphids and slowly shifts over to mulberry mealy-
bugs later (Jayaraj 2006). Sidde Gowda and 63.1.3 Biological Control
Kumar (1995) recommended Hibiscus cannabi-
nus as trap crop. Mealy bug population was sig- In West Bengal, India it is recommended to
nicantly low in mulberry with the trap crop release predatory ladybird beetles, Cryptolaemus
(3.14 %) compared to mulberry without the trap montrouzieri Mulsant @250 adults/ac or Scymnus
crop (11.44 %). As the trap crop facilitates better coccivora Ayyar @500 adults/ac in two split
colonization of M. hirsutus, it can also pave way releases during Oct-Nov and Jan-Feb to suppress
for preventing migration of the recommended the mealybugs (Santha Kumar et al. 1995;
predatory beetles from the release sites so that Anonymous 2010a).
they can effectively suppress the population of Complete control of M. hirsutus was achieved
M. hirsutus on mulberry. Manjunath et al. (2003) in Egypt, by introducing Anagyrus kamali Moursi
also indicated a signicant difference in mealy- (Encyrtidae) from Java, and then later in
bug damage in mulberry with H. cannabinus Caribbean islands and Florida. M. hirsutus
(4.28 %) as trap crop compared to mulberry as appeared on mulberry in California in 1999.
sole crop (26.02 %). Subsequently, the parasitoids Anagyrus kamali,
Samuthiravelu et al. (2005) reported that Gyranusoidea indica Shafee, Alam & Agarwal
mealybug infestation was minimized by reduced (Encyrtidae) and Allotropa sp. nr. mecrida
application of nitrogenous fertilizer blended with (Walker) (Platygastridae) were released for per-
neem cake @60 kg/ac (3 %) followed by pon- manent establishment on mulberry trees. The
gamia cake (4 %), mahua cake (4.4 %) and castor population density of M. hirsutus within the rst
cake (7.9 %) compared to control with recom- year was reduced by approximately 95 %.
mended dose of chemical fertilizer (20.8 %). Anagyrus kamali was the predominant parasitoid
Lavanya Latha et al. (2004) also reported that of M. hirsutus (Roltsch et al. 2006). Such intro-
limited irrigation once in 10 days and 25 % duction of A. kamali to India should be tried
reduced nitrogenous fertilizer applied in two split against M. hirsutus in mulberry gardens.
doses brought down the mealybug incidence to
1.60 % from 8.5 % in control with recommended 63.1.3.1 Integrated Pest
dose of fertilizer. In addition, Narendra Kumar Management (IPM)
et al. (2006) found that mealybug incidence was The Integrated Pest Management package against
more when nitrogenous fertilizer was applied as a pink mealybug includes clipping and destruction
63 Mulberry 583
of affected terminal portion, spraying of 0.2 % assumed the status of a major pest resulting in
DDVP with 0.5 % soap solution and release of C. huge losses to farmers in Tamil Nadu, Karnataka,
montrouzieri @250 adults/acre. The per cent Kerala (Shekhar and Qadri 2009; Krishnakumar
reduction in mealybug damage ranged from and Rajan 2009; Mahalingam et al. 2010).
73.21 to 88.81 whereas the increase in leaf yield
ranged from 3416.68 to 4750 kg/ha/year.
Narendra Kumar et al. (2006) also recommended 63.2.1 Damage
that the IPM practice involving the application of
5 % Neem seed kernel extract on 10th and 20th In mulberry, the papaya mealybugs infest leaf
day after pruning (DAP) integrated with release buds, leaves, stem portion, stump portion after
of predatory ladybird beetles @ 250/acre and top pruning, etc. They are found congregating all
clipping on 45th DAP proved better in control- along the veins on the lower side of the leaves.
ling the mealybug wherein the pest suppression Since they suck the plant sap continuously,
was recorded to an extent of 82.17 % (Manjunath affected leaves turn yellow and the plant growth
and Katiyar (1995). retards. In addition to sucking of plant sap they
also inject toxic substance through their saliva,
which causes deformation of plant parts. Due to
63.2 Papaya mealybug, profuse honey dew secretion, black sooty mould
Paracoccus marginatus secretion is also formed. When the mealybugs
infest with heavy population, the plants will end
Paracoccus marginatus popularly known as up with drying and death. Due to large quantity
papaya mealybug (PMB) has been accidentally of honey dew secretion, lots of ants will be
introduced in to south India and posing serious attending to them which arrive to feed on the
threat to several crops including mulberry. It sweet honey dew (Shekhar and Qadri 2009).
2011). However the indigenous predators are not tation was recorded (Qadri et al. 2011).
so effective in managing the huge populations of Surprisingly the pest incidence was reduced to
papaya mealybug. Since P. marginatus is an mere 1 % within 56 months of release. Saving
exotic pest, a classical biological control pro- the mulberry crop thereby increasing the cocoon
gramme was initiated, and the parasitoid A. papa- production has resulted in savings to the tune of
yae was imported by National Bureau of few crores of rupees in Karnataka.
Agriculturally Important Insects (ICAR),
Bangalore during July 2010 (Shylesha et al. Kerala Paracoccus marginatus appeared on
2010). The parasitoid was multiplied and released mulberry in 2009 in Idukki, Wyanad, Palakkad,
in farmers gardens through extension units of Malappuram, Thrissur districts of Kerala
Department of Sericulture of the southern states (Krishnakumar and Rajan 2009). Mulberry is
of India (Qadri et al. 2011). cultivated in about 300 acres in Kerala. Due to
release of Acerophagus papayae in 2011, mul-
berry crop worth few lakhs was saved. The suc-
63.3 Impact Analysis of Classical cess of classical biological control using A.
Biological Control of Papaya papayae has emerged as an excellent model
Mealybug in Mulberry reviving the sericulture to normalcy in the entire
in South India Tamil Nadu, Karnataka and Kerala.
Tamil Nadu There was 60 % damage by 63.4 Root mealybug Paraputo sp.
papaya mealybug in Tamil Nadu (T.N.). A total
area of 10,000 acres of mulberry gardens was Mulberry plantations in hilly areas of Northern
found infested with P. marginatus. It was esti- India such as Darjeeling and Kalimpong are
mated that mulberry crop worth Rs. 135 crores being infested by root mealybug, Paraputo sp.
was lost due the papaya mealybug infestation in (Pseudococcidae: Hemiptera) causing consider-
T.N. According to Qadri et al. (2011), more than able damage. It is considered as most persistent
33,000 adults of A. papayae were released (from and noxious pest (Biswas et al. 2002; Das et al.
Nov 2010 to March 2011) in the papaya mealybug 2004; Mukhopadhyay et al. 2010). It occurs
infested mulberry gardens of 350 farmers in the throughout the year with a peak during July-
districts of Erode, Tiruppur and Salem. After the August, and the population decreases with fall in
release of the parasitoids, the mealybug infestation temperature during winter months (Biswas et al.
was reduced from 90 % to less than 5 % thereby 2002; Anonymous 2011). It is a noxious pest
achieving a suppression of 8595 %. Similar con- which remains in the root zone as well as adja-
trol was achieved with the parasitoid in Trichy and cent to stump portion below the soil surface up to
in Coimbatore districts in Tamil Nadu. 20 cm or 3 deep and causes damage to root sys-
tem by sucking the sap (Biswas et al. 2002;
Karnataka A total of 15,000 adults of A. papa- Mukhopadhyay et al. 2010; Anonymous 2011).
yae were released (from Nov 2010 to Jan 2011) The affected mulberry becomes yellow and
in papaya mealybug infested mulberry gardens of stunted in growth (Misra et al. 1996).
150 farmers covering about 300 acres mulberry The mealybug causes appreciable damage to
in Chamarajanagar district. Further, a total of mulberry directly by sucking the sap and indi-
20,000 parasitoids were released in Mysore dis- rectly by making way for some fungal infection,
trict covering about 400 acres under seven leading to rotting of the root and ultimately death
Technical Service Centres (from Feb 2011 to of the plants. The infested mulberry plants show
May 2011). After the release of the parasitoids, vulnerability to the attack of various fungal
9095 % suppression in papaya mealybug infes- pathogens such as Fusarium solani, Phomopsis
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Research and Training Institute, Mysore, 1820 Karuppuchamy P, Samiayyapan R, Jonanthan ER
November, 2000 (2010) Invasive mealybugs of Tamil Nadu and their
Savithramma P, Dandin SB (2000) Leaf quality evaluation management. Karnataka J Agric Sci 23:69
of mulberry genotypes through chemical analysis. Williams DJ (2004) Mealybugs of southern Asia. The
Indian J Seric 39(2):117121 Natural History Museum/Southdene SDN. BHD,
Shekhar MA, Qadri SMH (2009) Papaya mealybug A Kuala Lumpur/London, 896 p
new menace to Mulberry in Tamil Nadu. Indian Silk Zaman A, Qader MA, Islam S, Barman AC, Alam MS,
48(4):2223 Islam M (1996) Effects of feeding of Tukra affected
Shekhar MA, Narendra Kumar JB, Sreenivas BT, Divya mulberry leaves on economic characters of silkworm,
SH (2011) Papaya mealybug, Paracoccus marginatus Bombyx mori L. Pak J Zool 28(2):169171
Tobacco
64
M. Mani and G.N. Rao
64.1 Species noticed during the crop season when hot weather
condition prevailed and rains were delayed.
Mealybugs are found to be injurious to tobacco About 2028 mealybugs were observed on the
(Nicotiana tabacum) in India, Zimbabwe, Africa, ventral side of 45 lower leaves of 1015 %
Italy, Argentina, etc. (Table 64.1). Phenacoccous plants. Crinkling of the lower leaves and pucker-
solenopsis (Tinsley) has been reported both in the ing in young leaves was observed (photo) due to
nursery and elds in India (Rao 2009; Bhatt the damage of the pest in Andhra Pradesh (Rao
2010). Heavy infestation of P. solani has been 2009). In Gujarat, P.solenopsis has been reported
reported to be found in Zimbabwe. as the major species. At the initial stage, the
mealybugs attach themselves to the lower leaves
and suck the cell sap. The infested leaves of
64.2 Damage tobacco showed sickly appearance, dried out
before maturity, and the quality of leaf also dete-
P. solenopsis appears in early sown tobacco nurs- riorated (Bhatt 2010).
eries and multiplies in large number and causes
damage to young leaves by sucking sap from the
succulent leaves. The affected leaves show puck- 64.3 Management
ering symptoms and become brittle during the
later course of development. As many as 19 Biological Control The Australian ladybird
mealybugs were recorded in each nursery bed. In beetle Cryptolaemus montrouzieri (Mulsant)
the main eld, mealybug damage was also (23 per tobacco plant) gave good control of
observed. The mealybugs were found on the ven- F. virgata in the glasshouse. The mealybug popu-
tral side of the lower leaves, and they were found lation declined from 16/cm2 to 0 after 35 days of
to suck the sap. Ants were also noticed visiting release (Gautam et al. 1988). C. montrouzieri can
the mealybugs for honeydew. This pest was also be used to control P. solenopsis on tobacco
(Rao 2009). In Gujarat, the encyrtid Aenasius
M. Mani (*) bambawalei (Hayat) was found on P.solenopsis
Indian Institute of Horticultural Research, (up to 30 % parasitism). Parasitized mealybugs
Bangalore 560089, India turned reddish brown, loss of white mealy pow-
e-mail: mmani1949@yahoo.co.in der from their mummied body (Bhatt 2010).
G.N. Rao
Central Tobacco Research Institute,
Rajamundry, AP, India
Chemical Since tobacco is a high-value crop, Bhatt NA, Jyani DB, Patel AD (2009) Mealybug
the leaf is used for human consumption; care is to Phenacoccus solenopsis Tinsley- An emerging pest of
bidi tobacco in Gujarat. In: Proceedings of national
be taken to select the chemicals for the control of symposium IPM strategies to combat emerging pests
mealybugs. Chloripyriphos 0.05 % spray gave in the current scenario of climate change, 2830 Jan
100 % control of the mealybugs in Andhra 2009. Chfcau,Pasighat, Arunachal Pradesh, p 38
Pradesh (Rao 2009). On tobacco, methomyl Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
90.80 % and profenophos had signicantly Pseudococcidae and Putoidae) with data on geograph-
reduced the mealybug population of P.solenopsis ical distribution, host plants, biology and economic
in Gujrat (Bhatt et al. 2009). importance. Intercept Limited, Andover, 686 p
Gautam RD, Paul AVN, Srivastava KP (1988) Preliminary
studies on Cryptolaemus montrouzieri Muls. against
the white tailed mealybug Ferrisia virgata (Cockerell)
References infesting tobacco plants. J Biol Control 82:1213
Rao GN (2009) Status paper on mealy bug on tobacco. In:
Bhatt NA (2010) Mealybug [Phenacoccus solenopsis Proceedings of interactive meeting on mealybugs and
Tinsley (Homoptera: Pseudococcidae)] a serious its management held at IIHR, Bangalore, 56th
pest of tobacco in Gujarat. Insect Environ December 2009, 146 p
16(2):9091
Jatropha
65
M. Mani
Biodiesel produced from nonfood crops like of jatropha were found infested with the mealy-
Jatropha (Jatropha curcas) is one of the most bug Rhizoecus bicirculus (McKenzie) (Ben-Dov
promising solutions for tackling the growing car- 1994). A prediction model has been developed,
bon emissions from transport. Paracoccus mar- which could act as an indicator of the severity of
ginatus (Williams and Granara de Willink) was the mealybug Paracoccus marginatus damage in
found to cause serious damage on jatropha in jatropha plantations, under tropical conditions, if
India (Regupathy, and Ayyasamy, 2009; Pretheep- no proper pest management measures had been
Kumar et al. 2013), Malaysia (Mastoi et al. employed (Pretheep-Kumar et al. 2013). The
2011), and Sri Lanka (Galanihe et al. 2010) The model for predicting the percentage of mealybug
infestation resulted in symptoms like crinkling or infestation in jatropha was of the form:
twisting of leaves and shoots, bunched and y = ax1b + cx2d, where y is the percentage of mealy-
unopened leaves, yellowing of leaves or leaf bug infestation, x1 is the mean monthly tempera-
drop, fruit drop, appearance of honeydew on ture, x2 is the mean monthly rainfall, and a, b, c,
leaves, sooty mould development, stunted d are the coefcients: a = 1.172; b = 1.951;
growth, deformation, and death of the plants in c = 3.722; d = 9.024. Standard error = 7.231;
case of severe infestation. Ferrisia virgata Correlation coefcient = 0.966. It was apparent
(Cockerell), Phenacoccus herreni (Cox and that the percentage of mealybug damage in jatro-
Williams), and Planococcus minor (Maskell) are pha decreased with increase in rainfall and vice
known to attack Jatropha sp. In California, roots versa (Pretheep-Kumar et al. 2013).
M. Mani (*)
Indian Institute of Horticultural Research,
Bangalore 560089, India
e-mail: mmani1949@yahoo.co.in
Ten natural enemies, including parasitoids have to be sprayed repeatedly to achieve satisfac-
viz. Acerophagus papayae (Noyes and Schauff), tory control. The combination of parathion and
Anagyrus loecki (Noyes), Pseudleptomastix mex- malathion with white oils makes spraying more
icana (Noyes and Schauff), and predators like efcient. To manage the insects at the beginning
Spalgis epeus (Westwood), Cryptolaemus of a local outbreak, severely infested branches
montrouzieri (Mulsant), Brumoides suturalis should be cut and burnt immediately (le:///C:/
(Fabricius), Cheilomenes sexmaculata (Fabricius), Documents%20and%20Settings/
Scymnus coccivora (Ayyar), Chilocorus sp., and user/y%20Documents/Downloads/Jatropha%
Chrysoperla zastrowi (Sillemi) (Esben-Petersen) 20under%20attack.pdf). The systemic acephate
were found attacking P. marginatus in India. on the plant can be used to clear up the mealy-
Among them, Acerophagus papayae was found to bugs on jatropha. Spraying is to be done twice at
be highly effective in controlling the mealybug 10-day intervals. Sprays can be scheduled in the
population in Bangalore North. early morning or evening when the temperatures
Mealybugs Ferrisia virgata and Planococcus are low. (http://articles.sun-sentinel.com/2000-
sp. suck the plants sap, resulting in yellowing, 06- 23/lifestyle/0006220403_1_mealybug-
withering and drying of plants, and shedding of toads-seeds)
leaves and fruits. The foliage and fruits become
covered with large quantities of sticky honeydew,
which serves as a medium for the growth of black References
sooty moulds, resulting in the reduction of the
photosynthetic area. Some ladybird beetles, Ben-Dov Y (1994) A systematic catalogue of the mealy-
bugs of the world (Insecta: Homoptera: Coccoidea:
including Cryptolaemus montrouzieri, Olla
Pseudococcidae and Putoidae) with data on geograph-
v-nigrum, and Azya luteipes, together with syr- ical distribution, host plants, biology and economic
phids such as Alloagrapta oblique, are known importance. Intercept Limited, Andover, 686 p
predators of mealybugs. Chemicals such as diazi- Galanihe LD, Jayasundera MUP, Vithana A,
Asselaarachchi N, Watson GW (2010) Occurrence,
non, malathion, dimethoate, and parathion are
distribution and control of papaya mealybug,
effective in controlling F. virgata. However, they Paracoccus marginatus (Hemiptera: Pseudococcidae),
65 Jatropha 593
an invasive alien pest in Sri Lanka. Trop Agric Res Ext (Jatropha curcas L.) based on the weather parameters.
13(3):8186 Int J Agrisci 3(6):440443
Mastoi MI, Azura AN, Muhammad R, Idris AB, Ibrahim Regupathy A, Ayyasamy R (2009) Need for generating base-
Y (2011) First report of papaya mealybug Paracoccus line data for monitoring insecticide resistance in new
marginatus (Hemiptera: Pseudococcidae) from invasive mealybug Paracoccus marginatus Williams and
Malaysia. Aust J Basic Appl Sci 5(7):12471250 Granara de Willink (Insecta: Hemiptera: Pseudococcidae),
Pretheep-Kumar P, Tilak M, Durairasu P (2013) A model the key pest of papaya and biofuel crop, Jatropha curcas.
for predicting the infestation of mealybugsin jatropha Resistant Pest Manag Newsl 19(1):3740
Forage Crops and Grasses
66
Narendra S. Kulkarni and M. Mani
Fodder crops and grasses harbour large number the infested areas, the plant stand is sparse and
of mealybugs throughout the world (Table 66.1). existing plants yield poorly and weeds often over-
Though a number of mealybugs are recorded on take these areas. The mealybugs produce white
grasses and fodder crops, only some are known to webbing and clusters of whitish eggs, so theyre
cause economic damage. often obvious in the soil. Ground mealybug is
restricted to the heavier soils. The eggs, nymphs
and adults all occur in the soil. Infestations in
66.1 Rhizoecus kondonis alfalfa elds generally occur in circular patches
and spread slowly. The damage to alfalfa plants is
The mealybug Rhizoecus kondonis Kuwana feeds very apparent in the summer months but less so
on alfalfa roots causing severe damage to alfalfa. It during the winter and spring (McKenzie 1967).
sucks out plant juices, which causes stunting and There are three generations per year. Mealybugs
yellowing of plants. The infestations generally start are abundant in July-August, December-January
in small circular areas near the eld borders and and March-April. Signicantly more R. kondonis
gradually increase in size up to an acre or so. Within were found 15.245.7 cm deep in the soil.
Table 66.1 List of mealybugs attacking the grasses and fodder crops
Mealybug species Plant species Region References
Antonina graminis (Maskell) Cyprus, Cyanodon, Many countries Ben-Dov (1994)
Echinochloa
Antonina martima Green Cyprus, Cyanodon India Williams (2004)
Sri Lanka
Antonina purpurea Signoret Grasses France, Italy & Spain Ben-Dov (1994)
Antonina graminis, A. indica Grasses Africa Williams (2001)
Hall., A. natalensis Brain &
A. transvaalensis Brain
Antonina graminis (Maskell) Bermuda grass, Brazil Culik and Gullan (2005)
Balanococcus botulus Cox Cyprus New Zealand Cox (1987)
Balanococcus poae Pasture grass New Zealand Charles et al. (2009)
(Maskell)
Balanococcus mediterraneus Cynodon Greece Kozar (1983)
Lozar
Balanococcus Cynodon Italy & Korea Ben-Dov (1994)
notodanthoniae Cox
Brevennia cyanadontis Cynodon & Sorgum Iraq Ben-Dov (1994)
(Bodennhemer)
Brevennia licus (DeLotto) Sorgum South Africa De Lotto (1967)
Brevennia rehi (Lindinger) Sorgum India David and Ananthakrishnan (2004)
Cyprus & grasses California Miller (1973)
Cynodon dactylon Australia & Papua New Williams et al. (1981)
Guinea
Chaetococcus australis Cyperus Australia Ben-Dov (1994)
(Froggatt)
Chlorozococcus sorghi Sorgum India Williams (2004)
Williams
Dysmicoccus andropogonisn Andropogon grass India Williams (2004)
sp.n.
Dysmicoccus boninsis Sorghum & Cynodon Ben-Dov (1994)
(Kuwana)
N.S. Kulkarni and M. Mani
Mealybug species Plant species Region References
Dysmicoccus brevipes Maize & Cyprus Ben-Dov (1994)
(Cockrell) & Dysmicoccus
neobrevipes Beardsley
Dysmicoccus multivorus Lucerne Turkmenia & USSR Myartseva and Kharchenko (1988)
Koteja & Zak-Ogaza
Ehrhornia cupressi (Ehrhorn) Cyprus California & Mexico Ben-Dov (1994)
Fonscolombia butorinae Grasses Russia Danzig (2007)
(Danzig et Gavrilov)
Formicoccus lingnani Sorghum Malaysia Williams (2004)
66 Forage Crops and Grasses
(Ferris)
Ferrisia virgata (Ckll.) Su-babul Leucaena India Pillai and Gopi (1990)
leucocephala
Maize & Lucerne Ben-Dov (1994)
Geococcus coffeae Green Cyperus Many countries Ben-Dov (1994)
Heliococcus singularis Cyperus India Williams (2004)
Awasthi & Shafee
Heterococcus cyperi (Hall) Cyperus Egypt Ben-Dov (1994)
Marendelleae harrisae Sorhgum Nigeria Ben-Dov (1994)
Williams Leucaena http://www.plantwise.org/KnowledgeBank/Datasheet.aspx?dsid=36335
Paradoxococcus mdaniell Sorghum Texas Ben-Dov (1994)
McKenzie
Phenacoccus angustatus Sorghum Kazakhstan Ben-Dov (1994)
Borchsenius
Phenacoccus bicerarius Sorghum Kazakhstan Ben-Dov (1994)
Borchsenius Colver Europe
Phenacoccus hordei Grasses England Malumphy (2011)
(Lindeman)
Planococcoides lindingeri Sorghum Egypt & Israel Ben-Dov (1994)
(Bodenheimer)
Planococcus minor (Maskell) Maize Ben-Dov (1994)
(continued)
597
Table 66.1 (continued)
598
Rhodesgrass mealybug was an important pest occurring at only 20 % of survey sites. In addi-
of forage and lawn grasses in Texas. Cultural tion, N. sangwani exhibited a patchy geographic
control includes collecting and destroying grass distribution. Possible causes for these results are
clippings. A classical biological control project that N. sangwani has not dispersed widely since
resulted in the introduction of several species of its introduction, or that the imported re ant,
parasitoids (Schuster et al. 1971). However, Solenopsis invicta Buren, is interfering with bio-
complete control was attained by encyrtid para- logical control. Two other encyrtid wasps
sitoid Neodusmetia sangwani (Rao), imported Acerophagus sp. and Pseudectroma sp. are utiliz-
from India after being disseminated by aircraft ing A. graminis as a host (Chantos et al. 2009.
(Schuster et al. 1971: Dahlsten and Hall 1999).
By 1976, it was estimated that the parasitoid
saved ca. 17 million dollars annually in turf grass 66.5 Dysmicoccus brevipes
management costs (Dean et al. 1979). The exotic
parasitoid Anagyrus antoninae Timb. established Dysmicoccus brevipes (Cockerell) is a polypha-
and gave good control in cooler and more humid gous mealybug; it was only found in moderate
areas of Texas, Mexico and Florida. Antonina densities on Rhodesgrass, Chloris gayana, and
graminis is resurging as an important pest of turf- wire grass, Eleusine indica, both of which were
grass across Texas and the South-eastern United found in mowed and unmowed weedy areas with
States. This mealybug is known to feed on many the former species being more common in
warm-season turf grasses and pasture grasses. Hawaii. All phenological stages of Rhodesgrass
Cultivars of kikuyugrass (Pennisetum clandesti- were infested with pink pineapple mealybugs,
num Hochst) and bermudagrass (Cynodon spp.) but only mature wire grass plants were infested
were signicantly more susceptible than cultivars (Gahan) (Pandey and Johnson 2006).
of seven other genera of turfgrass. Cultivars of St.
Augustinegrass, buffalograss and zoysiagrass
each exhibited susceptibility of >2 mealybugs 66.6 Brevennia rehi
per 7.5 7.5-cm plant. Populations did not
exceed < =0.5 mealybug per plant on centipede- Many grass species are grown as lawns or used in
grass, seashore paspalum, bahiagrass, or tall fes- golf courses and other recreational settings espe-
cue (Reinert and Vinson 2010). A survey cially turf grasses are susceptible to numerous
conducted in south-eastern United States indi- species of pest insects, and annually millions of
cated that N. sangwani was uncommon overall, dollars are spent to prevent or eliminate infesta-
66 Forage Crops and Grasses 601
tions. The Tuttle mealybug, Brevennia rehi Bermuda and zoysia are important lawn grasses,
(Lindinger), is a pest of many grass species and especially in the southern United States, infesta-
occurs nearly worldwide, especially where rice tion by Tuttle mealybug should be considered
and sugarcane are grown. Ofcially recorded in whenever dieback is noticed, especially if the
the United States only from Arizona, California, grass blades show white wax or are sticky from
Florida and Texas (Ben-Dov 2012), but is proba- honeydew secretion. Both Bermuda and zoysia
bly more widespread in south-eastern states that lawns are commonly installed as sod or plugs,
produce turf grasses for the sod market. Tuttle which provide a ready route for the spread of
mealybug is known from other regions of the infestations should the pest control practices of
world (e.g., Palearctic region Afghanistan, Iraq, the grower fail to maintain a pest-free production
Iran; Ben-Dov 2012) where there is a distinct environment.
cold season, which suggests the mealybug may
be capable of surviving in much of the United
States. Tuttle mealybug was described in the 66.7 Trionymus winnemucae &
United States as Heterococcus tuttlei Miller and Saccharicoccus sacchari
its taxonomy status is now amended to junior
synonym of Brevennia rehi. Healthy turfgrass Trionymus winnemucae McKenzie (Winnemuca
will have lower mealybug populations, so proper grass mealybug) lives within the sheath, but may
fertilization and watering is needed. Keep bene- also be found below the crown at the crownsoil
cial insects in the area to reduce the number of interface. Saccharicoccus sacchari (Comstock)
mealybugs, such as big-eyed bugs and lady bee- (pink sugarcane mealybug) is also an elongated
tles. After mowing, collect and destroy all pinkish mealybug that will occasionally infest
infested grass clippings. Brevennia rehi is also ornamental grasses. Both T. winnemucae and Sa.
recorded from Israel, Iraq, Azerbaijan, and sacchari are slightly larger, with a more elongate
Tajikistan and from Brazil. The rice mealybug is body form.
recorded from Israel as a pest to lawn grasses,
Dactyloctenium australe (Poaceae) (Ben-Dov
2008). Mealybugs hide between the grass blade 66.8 Balanococcus poae
and the stem where they can be difcult to see.
They are destructive infestations in South Asia The pasture mealybug Balanococcus poae
(e.g., India, Bangladesh). There was a correlation (Maskell) can be a serious pasture pest in
between drought stress and degree of infestation, Canterbury areas, and is known to occur in
possibly due to an increase in the availability of Manawatu and Nelson region of New Zealand.
amino acids in the vascular uid (Dale 1994). It Adult mealybugs are small, growing to about
was discovered infesting bermudagrass (Cynodon 2 mm in size, pink in colour. There are white,
dactylon) seed production crops in Arizona to waxy secretions in the plant crown and upper
such an extent that the sticky exudates produced roots. These will appear as cotton-woollike glob-
by the mealybugs fouled the harvesting equip- ules. Damage becomes apparent in autumn during
ment (Miller and McKenzie 1970). Tuttle mealy- extended dry periods. Symptoms of infestation
bug is a recorded host of the parasitoids Rhopus resemble those of drought, with pastures brown-
nigroclavatus Ashmead and Apoleptomastix ing off. The mealybug damage tends to affect a
bicoloricornis Girault (Hymenoptera: whole paddock, rather than isolated patches. It
Encyrtidae) (Noyes 1988). Rhopus nigroclavatus causes widespread ryegrass death, leading to poor
does not occur in Florida, but is recorded from pasture persistence. At Hawke's Bay, adult
several other states, and A. bicoloricornis is not females of Balanococcus poae were found
recorded from the United States. Because throughout the year, typically in wax cells ca.
602 N.S. Kulkarni and M. Mani
12 cm below the soil surface, with a peak density The timing of life-stages indicates that there was a
of ca. 1,300/m2 during winter and early spring single generation each year, but a partial second
(June-October). Winter eggs were followed by generation may also have occurred in late sum-
neonate nymphs from spring through summer. mer. No males were found (Charles et al. 2009).
Pasture mealybug (Balanococcus poae) was high number on any individual plant and the
found infesting native grasses, tussocks in symptoms of infestation on the surface become
Poaceae and several introduced pastoral grass easily visible. Usually, the mealybug is capable
species particularly ryegrass (Lolium spp.) in of growing to a maximum length of 4 mm (3/16
Canterbury, New Zealand. Pasture mealybug are in.) and it thrives in the narrow gap between stem
capable of inicting severe damage to endophyte- and the enfolding leaf sheath. Initially, this pest
free ryegrass (Pennell et al. 2005). The use of generally forms colonies near the base of the
pure stands of endophyte-infected grasses or a plant and gradually ascends upward with the
mixed stand of infected and non-infected plants amplication of their population. In general, the
may increase the persistence and durability of initial surface symptoms of a plant being infested
turf and forage grass species in the presence of by Miscanthus mealybug include slowing down
foliar damaging mealybugs (Sabzalian et al. of the plant's growth and an abnormal entwining
2004). of the owering head. In addition, the colour of
the sheath tissue as well as the stem becomes
deep red in parts where the mealy bugs are draw-
66.9 Miscanthicoccus miscanthi ing their food from the plant, particularly in the
later part of the growing season. Even when a
Miscanthus is a genus of about 15 species of plant is severely plagued, it is not eliminated, but
perennial grasses native to subtropical and tropi- is decreased to ugly, distorted masses as the white
cal regions of Africa and southern Asia. The dis- powdery wax swathes its stems, particularly in
persal of the Miscanthus mealybug, the lower parts. Plants that are infested by this
Miscanthicoccus miscanthi (Takahashi), has bug usually become incapable of owering in any
actually been unsuspectingly rapid, by means of way. In some cases, the owering stalks may pos-
selling as well as exchanging the plants infested sibly be underdeveloped resulting in the owers
with this insect. It only becomes conspicuous to open droopingly among the foliage, instead of
when the population of this bug reaches a very blossoming elegantly above.
66 Forage Crops and Grasses 603
enclosing the pistillate spikelets. Injury appeared pronounced that the species is supposed to be
as foliar yellowing with the most severely injured extinct locally (Brinon et al. 2004).
plants turning straw-brown and dying (Baxendale
et al. 1994). Pubescent leaves increase buffalo-
grass susceptibility to mealybugs Tridiscus spo- 66.18 Sorghum
roboli and Trionymus sp. (Johnson-Cicalese et al.
1998, 2011). Parasitism on these mealybugs by Pink sugarcane mealybug Saccharicoccus sac-
Rhopus nigroclavatus in Nebraska went up to chari (Cockerell) sucks the sap and reduces cane
78.5 % indicating its potential in the population vigour, besides causing sooty mould. The mealy-
regulation of these mealybugs (Heng-Moss et al. bugs are usually attended by ants. Severe attack
1999). results in stunted growth, yellowing of leaves,
deposition of sticky honeydew, and development
of sooty mould on the mealybug infested plants.
66.15 Pseudococcus saccharicola
Takahashi
66.19 Maize
It was found infesting Chloris barbata Sw. (swol-
len ngergrass), C. radiata (L.) (radiate nger- Infestations of Ferrisia viragata remain clustered
grass) and Cynodon dactylon L. (bermudagrass) around the terminal shoots and leaves, sucking
in Guana Island, and nearby Beef Island and the sap which results in yellowing, withering and
Tortola, in the British Virgin Islands (BVI). The drying of plants and shedding of leaves. The
coccinellid predator Hyperaspis scutifera sooty moulds and waxy deposits result in a reduc-
(Mulsant) was commonly witnessed with colo- tion of photosynthetic area.
nies of P. saccharicola on all three islands
(Wheeler et al. 2010).
66.20 Forage Trees
cards can be used to trap the ying adult males, duced in New Caledonia, Heliococcus summervillei
Brookes (Hemiptera, Pseudococcidae). [French]. Bull
preventing them from mating. Insecticidal soaps
Soc Entomol France 109(4):425428
and horticultural oils work great in controlling Chantos JM, Vinson SB, Helms KR (2009) Distribution
the mealybugs. The tricky part is mealybugs tend and abundance of parasites of the rhodesgrass mealy-
to hide very well where leaves attach to the stem, bug, Antonina graminis: reassessment of a classic
example of biological control in the southeastern
so make sure you get coverage there. Horticultural
United States. J Insect Sci (Madison) 9:48
soaps and oils dont have systemic properties, Charles JG, Chhagan A, Forgie SA, Slay MWA, Edwards
which means when spraying, the product must RD (2009) Observations on the biology of the pasture
come in contact with the pest. Burn leaves with mealybug, Balanococcus poae, from Hawke's Bay
pastures. NZ Plant Protect 62:197204
horticultural soaps and oils. These products need
Clausen CP (1978) Introduced parasites and predators of
to be applied when the air temperature is cool. arthropod pests and weeds: a world review. USDA
Make sure your plants were watered well the day ARS Agric. Hand Book, University of Illinois No
before applying any control measures. Following 480545 p
Cox JM (1987) Pseudococcidae (Insecta: Hemiptera).
labelled rates also reduces the risk of leaf dam-
Fauna NZ 11:1228
age. More is not better. Also, make sure bene- Culik MP, Gullan PJ (2005) A new pest of tomato and
cial insects are not affected while spraying other records of mealybugs (Hemiptera:
insecticides. There are a few benecial insects Pseudococcidae) from Espirito Santo, Brazil
(Summary In Portuguese). Zootaxa 964:18
that can help in mealybug treatment, too. Green
Dahlsten DL, Hall RW (1999) Biological control of
lacewings feed on the crawler stage of almost any insects in outdoor urban environments. In: Bellows
mealybug, where some others are more special- TS, Fisher TW (eds) Handbook of Biological Control:
ized like the mealybug destroyer (Cryptolaemus Principles and Applications. Academic Press, San
Diego/New York, 1046 p
montrouzieri). This benecial insect is a type of
Dale D (1994) Insect pests of the rice plant-their biology
ladybug that loves to feed on most mealybug spe- and ecology. In: Heinrichs EA (ed) Biology and man-
cies. Mealybugs can be controlled if the timing of agement of rice insects. Wiley Eastern Limited, New
the initiation of the treatment is planned correctly Age International Limited, Bangalore, pp 363486
Danzig M (2007) Mealybugs of the genus Fonscolombia
at the crawler stage.
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Forest Plants
67
R. Sundararaj and M. Mani
Mealybugs often cause serious damage to the highly polyphagous and have many collateral
growth of forest tree species particularly in nurs- hosts and hence, they can spread very rapidly to
eries and plantations. The damage is caused by the neighbouring plants.
sap sucking resulting in dieback symptoms and
secreting copious amount of honeydew on which
black sooty mould fungus develops. Often the 67.1 Ferrisia virgata
infestation results in drying of branches causing
dieback of branches and ultimately death in seed- The striped mealybug Ferrisia virgata (Cockerell)
lings and trees. The affected owers wither and is covered with powdery white wax and has a pair
fruits dry up, fall off prematurely. The seedlings of purplish dorsal stripes along the back. It is
and trees affected severely by mealybugs shed reported to breed on leaves, stem and fruits of
their leaves and look like sickly appearance and large number of tree plants including Azadirachta,
in some cases drying of branches in trees and Anacardium, Annona, Artocarpus, Caesalpinia,
death of seedlings. Most of the mealybugs are Casuarina, Cassia, etc. in India (Ali 1970). On a
R. Sundararaj (*)
Institute of Wood Science and Technology, M. Mani
Bangalore 560 003, India Indian Institute of Horticultural Research,
e-mail: rsundariwst@gmail.com Bangalore 560089, India
variety of host plants, it was most active during and owering. During this period, it infests
August-November and March-April but very almost all parts of tree. The affected owers
much reduced during December-January (Rawat wither and fruits dry and fall off prematurely. Its
and Modi 1968). Ferrisia virgata was found infestation starts from February reaching peak
infesting Santalum album in India (Sundararaj during March and April and then it declines in
et al. 2006). Karnataka (Mangala et al. 2012). An infestation
It is emerging as an important pest on by F. virgata was reported on four tree species
Pongamia pinnata. Nymphs and adults were (Albizia lebbeck, Gliricidia sepium, Leucaena
found sucking the sap from both the surfaces of leucocephala and Cassia siamea) in a screen
leaves as well as tender shoots and owers of P. house at IITA main station, Ibadan, Nigeria
pinnata at the time of formation of new foliage (Kadiata et al. 1992).
nicantly higher on branches of the woody an important role in maintaining N. viridis popu-
legume Leucaena leucocephala in northern lations at low levels (Nechols and Seibert 1985).
Guam, Mariana Is. Cryptolaemus was found
feeding on N. viridis (= N. vastator) infesting
several plants in Guam but the predator was not 67.3 Nipaecoccus lamentosus
feeding preying N. viridis when it occurred on
woody legume L. leucocephala. It might be due Nipaecoccus lamentosus (Cockerell)
to the presence of amino acid mimosin derivative (Pseudococcus lamentosus) has been recorded
which might have acted as feeding deterrent on limes, Tamarix stricta, Ficus carica [gs],
through the host (Muniappan et al. 1980). The Vitis sp. and Nerium oleander in Iran. There were
presence of the ants Technomyrmex albipes (Fr. four generations annually in the Fars region. The
Smith) decreased the percentage of N. viridis coccinellid Cryptolaemus montrouzieri has been
parasitized by the encyrtid Anagyrus indicus imported from northern Iran and has proved to be
Shafee et al. and the mortality of mealybugs was effective as a biological control agent of N. la-
attributed due to parasitism by A. indicus and pre- mentosus (Khalaf and Aberoomand 1989).
dation by other arthropods. Natural enemies play
M. hirsutus (Green) (PHMB) was known to infest Humococcus resinophilus (Green) in northern
Acacia arabica and Albizzia lebbak in Egypt. India is a pest of Pinus roxburghii regeneration.
Maconelicoccus hirsutus was detected in teak As a result of heavy infestations, branches appar-
plantations in 2004 in the Banderas valley in ently turn black and die (Ben-Dov 1994).
Mexico. A biological control programme was ini-
tiated in May 2004 to release 210,000 of the
predator Cryptolaemus montrouzieri on 150 ha. 67.8 Paracoccus marginatus
Damage to trees was reduced by 92 %. In India,
it has been reported on Samanea saman, Tectona In multi-tier agroforestry ecosystems of Kerala,
grandis, Tabeubuia rosea, Delonix regia (Anand India, the invasive mealybug Paracoccus margin-
Persad and Khan 2006), Ficus cunia, F. religiosa, atus Williams and Granara de Willink infestation
F. indica (Ayyar 1930; Varshney 1992). In Bahia was reported from teak, rubber, and other such
de Banderas, Nayarit, Mexico, the parasitoid plantations of Kerala even though the incidences
Anagyrus kamali Moursi regulated the popula- were highly localized. In the case of young teak
tion growth of Maconellicoccus hirsutus on teak. plantations, the immediate action taken was to
The average reduction of the pest was 96.5 % in chop off the infested branches and burn them.
30 days after release (Garcia-Valente et al. 2009). Subsequently the exotic parasitoid Acerophagus
M. hirsutus was found infesting Casuarina equi- papayae was released in the forest ecosystem to
setifolia in Andhra Pradesh (Murthy et al. 1997) control Pa. marginatus.
and Ficus trees in Gujarat, India (Muralidharan
and Badaya 2000).
67.9 Planococcus vovae
67.10 Phenacoccus azaleae but they may severely retard growth (http://for-
estpests.org/vd/7047.html). Three native parasit-
The Bunge Prickly-Ash tree plant (Zanthoxylum oids, Zarhopalus debarri Sun, Acerophagus
bungeanum) damaged by the mealybug coccois Smith and Allotropa oracellae Masner
Phenacoccus azaleae Kuwana which attracts its help regulate this mealybugs population size in
natural enemy, the ladybug Harmonia axyridis the southeast United States. All three parasitoids
(Pallas), was studied in Tainhang Mountain Area were imported to China and released in heavily
of Shanxi Province, China, during 19992001 infested slash pine plantations (Clarke et al. 2010).
(Xie et al. 2004).
The mealybug Oracella acuta (Lobdell) is native Judas tree, Cercis siliquastrum, is a small decidu-
to the south-eastern United States. Hosts of this ous tree from Southern Europe and Western Asia
mealybug include loblolly (Pinus taeda L.), slash which is noted for its prolic display of deep pink
(Pinus elliottii Engelm.), Virginia (Pinus virgin- owers in spring. Heavy predation by
iana Miller), shortleaf (Pinus echinata Miller), Cryptolaemus montrouzieri was observed on Ps.
and longleaf (Pinus palustris Miller) pine (http:// viburni (Signoret) infesting Judas trees in ave-
forestpests.org/vd/7047.html). O. acuta was acci- nues of Turin resulting in small mealybug popu-
dentally introduced into Guangdong, southern lation in subsequent years (Arzone 1983).
China, in 1988 on scions of slash pine (Pinus
elliottii) and found damaged pine trees (Sun
Jiang Hua et al. 1996). Mealybugs either settle on 67.11 Peliococcus serratus
the shoot or occasionally between the needles
near the fascicle. Females secrete a characteristic American Beech Fagus grandifolia is an impor-
white resin cell that covers their body. The tips of tant tree in forestry. Peliococcus serratus (Ferris)
new shoots are the preferred settling site, though was known to attack F. grandifolia in Maryland,
the entire shoot may be colonized when popula- USA. The mealybug had two generations in a
tions are high. The resin cells, shoots, and nee- year. The eggs were laid in an ovisac on the bark
dles may become covered with black, sooty in June-August (hatching in 710 days) and in
mould growing on honeydew produced by the October-November (these overwintering, hatch-
mealybug. Infestations rarely cause tree mortality, ing in late April or early May). Limiting factors
612 R. Sundararaj and M. Mani
Dysmicoccus obesus (Lobdell) was found in Serianthes nelsonii is a large tree endemic to
Arkansas living in crevices under bark scales of Guam and Rota of the Mariana Islands. Three
loblolly pine trees (Pinus taeda). Most individu- species of mealybugs, Dysmicoccus neobrevipes
als (77 %) were found on the bole between 0 and Beardsley, D. brevipes (Cockerell), and
90 cm of the ground, and they showed slight pref- Planococcus citri (Risso), feed on the leaves, leaf
erences for the northern and southern bole expo- buds, branch tips, and roots of trees and seed-
sures. Individuals of the formicid Crematogaster lings. On the cultivated tree in Yona, up to 40 %
were observed tending the mealybug. Three of the branch tips were killed every two weeks by
broods per year were detected, with adults pro- a combination of D. neobrevipes and P. citri.
duced in May, July and September. It is suggested Most mealybug colonies were removed by preda-
that D. obesus probably overwinters off the tree tors, including the lady beetle Nephus roepkei
as immatures. The documented occurrence of D. (Fluiter) (Coccinellidae). Seedlings may remain
obesus from ten southern and south-eastern states vulnerable to mealybugs for longer periods of
in the USA suggests that its distribution is prob- time; malathion effectively killed the mealybugs
ably throughout the range of its host, P. taeda. on the seedlings (Gary et al. 1996).
Records from Maryland indicated that the pseu-
dococcid also feeds on Virginia pine (P. virgin-
iana) (Thompson and Colvin 1990). 67.21 Management
Clausen CP (1978) Introduced parasites and predators of Lotfalizadeh H, Ahmadi AA (2000) Natural enemies of
arthropod pests and weeds: a world review. USDA cypress tree mealybug, Planococcus vovae (Nasonov),
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principles and applications. Academic, San Diego/ Pongamia pinnata (L.) Pierre and their population
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Galanihe LD, Watson GW (2012) Identication of 20(1):11015
Rastrococcus rubellus Williams (Hemiptera: Martorell LF (1940) Some notes on forest entomology.
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Garcia-Valente F, Ortega-Arenas LD, Gonzalez- (Homoptera: Pseudococcidae) in Greece: three new
Hernandez H, Villanueva-Jimenez JA, Lopez-Collado records. Hellenic Plant Protect J 1(1):3538
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Monthly Magazine 121:53 Bitki Koruma Bulteni 38:2341
Glasshouse, Greenhouse
and Polyhouse Crops 68
K. Gopalakrishna Pillai
Protected environments such as glasshouse/ feeds upon perennial crops such as cycad and
greenhouse/net house/polyhouse are those that Phormium tenax. Ferrisia virgata (Ckll.) appears
maintain plants year round. They provide optimal in a very severe form on poinsettia in the poly-
conditions for insect and mite pests to survive, house. In glasshouse, the obscure mealybug,
develop, and reproduce. Mealybugs are serious Pseudococcus viburni (Signoret), and
pests of various crops in greenhouses and proba- Phenacoccus gossypii (Townsend and Cockerell)
bly the most difcult-to-control pests in green- are found on chrysanthemum. Maconellicoccus
houses. Mealybugs are not particular about their hirsutus (Green), Paracoccus marginatus
hosts, and probably all species of crops are sus- (Williams and Granara de Willink), and
ceptible to mealybugs, especially when cultivated Phenacoccus solenopsis (Tinsley) are also found
in protected environments. on several crops in greenhouses. Phenacoccus
madeirensis (Green) has become an increasingly
damaging pest in greenhouse ornamental produc-
68.1 Mealybug Species tion. Maconellicoccus hirsutus is known to attack
many species of ornamental plants including
There are a number of different mealybugs of Allamanda, Angelica, Anthurium, Bougainvillea,
concern to greenhouse growers. In greenhouses Croton, ginger lily, Heliconia, Ixora, hibiscus,
of California, the most frequently found mealy- palm, and oleander. The lily bulb mealybug
bugs are the long-tailed mealybug, Pseudococcus Vryburgia amaryllidis (Bouch) and the obscure
longispinus (Targioni-Tozzetti) and the citrus mealybug Pseudococcus viburni were commonly
mealybug, Planococcus citri (Risso) (Lain and found as well. Vryburgia amaryllidis is limited to
Parrella 2004). Planococcus citri is the most a few plant families (especially Liliaceae and
common and damaging insect pest in greenhouse Iridaceae). It occurs on the bulb and on the basal
and protected cultures. With the exception of portion of the leaves. Pseudococcus viburni was
roses, P. citri feeds on many short-term crops found both on the roots and the aerial portion of
such as coleus, whereas P. longispinus often the plants, most commonly on short-term crops.
Root mealybugs (Rhizoecus spp.) feed on the
root systems of plants; so, they can be undetected
for long periods of time. Phenacoccus solenopsis
K.G. Pillai (*) occurs more commonly on the roots, stems, and
Indian Institute of Horticultural Research,
Bangalore 560089, India foliage close to the soil line in dry climates, com-
e-mail: pillaig@iihr.ernet.in pared to settling on the upper foliage of the plant.
The most common species of mealybugs that Monitoring of immature and adult mealybugs is
infest crops are immediately recognized in the to be carried out on the stems, leaves, and ow-
adult stage by the white, yellowish-white, ers. The mealybugs survive several millimeters
whitish-gray, or pale pink to pale blue color coat- below the soil surface. Observations should be
ing. Mealybugs can be found on all plant parts, directed to all plant tissues for the white waxy
especially roots, rhizomes, pseudobulbs, and the specimens. Sticky traps set out in the greenhouse
underside of leaves. They are adept at hiding on can be used to detect the presence of mealybug.
roots and rhizomes deep in the potting media, in Once mealybugs become established, it is dif-
crevices, and under sheaths. Mealybugs are cult to achieve effective control. Incoming plants
active and crawl from one plant to another, pot to should be inspected for signs of mealybugs.
pot, and across benches. Mealybugs hide under Roots of newly purchased plants should be
rims of pots and trays, in bench crevices, and inspected for the root mealybug. Greenhouses
even drop from overhead plants. The mealybugs should be kept as weed-free as possible.
species have the ability to increase rapidly in Operational parameters of traps baited with the
population size in a relatively short period of pheromones of three mealybug species were
time. With their piercingsucking mouthparts, optimized in nurseries producing ornamental
they feed on leaf and stem axils, and even on the plants. All pheromone doses (1320 g) attracted
roots of some plants. The mealybugs damage the P. longispinus and P. viburni males, with the low-
plant by extracting the sap, which stresses the est dose (1 g) attracting the fewest males for
plant, resulting in the leaves becoming chlorotic both species. Doses of 3.2100 g were as attrac-
and shedding over time, as well as fruit bodies tive to male P. longispinus as the highest dose
being aborted. Flowers often take on an abnormal (320 g); doses from 10 to 320 g were equally
shape, reducing yield. Infested leaves become attractive for P. viburni males. Lures containing
curled and crinkled, acquiring a rosette pattern, 25-g doses of either pheromone had effective
with the plant appearing bushy and stunted. In eld lifetimes of at least 12 weeks. When
addition, the high numbers of developing mealy- pheromone-baited traps for P. longispinus were
bugs produce large amounts of honeydew that compared with manual sampling, trap counts of
fall onto the lower leaves, producing a substrate male mealybugs were signicantly correlated
for the development of sooty mould, which inhib- with mealybugs counted on plants in the vicinity
its photosynthesis within the plant. of the traps (Waterworth et al. 2011).
Mealybugs can be serious and persistent pests
in the greenhouse. Host plant range depends on
the particular mealybug species, and includes 68.4 Management
herbaceous annuals or perennials, foliage plants,
orchids, vegetables, and herbs. Some of the Mealybug management in greenhouses is dif-
greenhouse crops prone to mealybug infestations cult because of their propensity to move into the
include coleus, croton, dracaena, hoya, English potting medium and feeding on roots, or for the
ivy, cus, fuchsia, stephanotis, schefera, hibis- crawlers to work their way into tight places.
cus, mandevilla, strawberry plant (houseplant), Repeated application of any treatment is required
jade plants, palms, prayer plants, gardenia, and to kill the immature, and treatments are at their
orchids as well as many other foliage plants. The greatest effectiveness against the small crawlers.
mealybugs have been found feeding on mari- All control efforts must begin immediately
golds, gerbera, daisies, poinsettias, begonias, and following discovery. Even light infestations
chrysanthemums. restricted to one or a few plants can explode
68 Glasshouse, Greenhouse and Polyhouse Crops 623
rapidly and necessitate chemical methods. When can live up to 6 weeks and can continue to repro-
required, infested plants should be isolated duce after crop harvest. Severe pruning of
immediately from others to prevent the mealy- infected plants can be considered to allow for
bugs from moving among them. Also, the lips better spray coverage, followed by an aggressive
and cracks of pots, trays, and benches should be control program. As soon as an infestation is
checked, because females will wander and leave detected, infested plants should be isolated and
the plant to nd hiding places. The physical con- treated. It is important to prune or cut infested
ditions in greenhouses approach an optimum stems or branches from plants and destroy the
environment for the uncontrolled increase in infested plant material. Also, stalks and crop resi-
populations of phytophagous insects. Once the due in infested sites should be removed and
insect is introduced, the greenhouse structure destroyed, as such residue left in the greenhouse
affords warm temperature, high humidity, and a can harbor mealybugs, which can survive to
physical barrier, isolating the pest from the natu- invade the new crop. It is necessary to sanitize
rally occurring predators and parasites. Effective equipments and check clothing items to prevent
chemical control of insects in greenhouse condi- the transfer of the pest into new locations. Small
tions when plant diversity is high is difcult. populations of mealybugs can be controlled by
Compact growth habit, certain structural plant inspection of plants, removing, and handpicking
forms such as leaf sheaths, and dense foliage, all the specimens from newly infested plants. Soap
prevent adequate application of the chemical to applications are often effective against targeted
the entire plant. Failure to treat all surfaces, along small populations of the mealybug.
with sublethal dosages due to improper applica-
tion rates and pest diversity contribute to a seri-
ous problem encountered by greenhouses 68.6 Chemical Control
todayresistance and resurgence. The phenom-
enon of pesticide resistance followed by a rapid The conventional management tactics for mealy-
resurgence of the surviving insects can cause an bugs in greenhouse ornamental production
actual increase in the pest population following include regular application of insecticides.
chemical application. Persistent populations of mealybugs or infesta-
tion in many plants may demand the need for use
of synthetic insecticides. Well-established infes-
68.5 Cultural/Physical/mechanical tations are difcult to control, because their waxy
control and sanitary secretions help to protect the young nymphs and
measures eggs from penetration with chemical sprays. The
crawler stage, which does not possess a waxy
Prevention is the most important element of covering, is most susceptible to insecticides,
mealybug control. Careful selection of clean cut- including insect growth regulators (e.g., azadi-
ting materials before propagation is critical. If rachtin, buprofezin, and kinoprene), insecticidal
needed, the cutting materials could be treated soaps (potassium salts of fatty acids), horticul-
with pesticides before rooting. Planting material tural oils (petroleum-based), and possibly insect-
should not be taken from infested elds. It should killing fungi (Beauveria bassiana).
be made sure that transplants are clean and The types of insecticide applications include
healthy before introducing into elds. Sanitation foliar sprays and those directed toward the grow-
is the second most important element of control. ing medium (drench or granule). Adult mealy-
Fields should be scouted regularly, checking bugs are difcult to manage, because they form a
entire plants, paying attention to ants and other white, waxy protective covering that is nearly
crawling insects that move mealybugs. impervious to most insecticides. And, because
Aggressive control programs, if present, could be most insecticides have no activity on eggs (with
implemented immediately. Sanitation in green- the possible exception of petroleum-based or
houses and shade houses is critical. The female neem oils), at least 23 weekly applications
624 K.G. Pillai
usually are required to achieve satisfactory sup- Growth Regulators and Chitin Inhibitors are
pression, especially when dealing with overlap- classes of insecticides that have some potential
ping generations. Although very few (if any) for mealybug management. The insect growth
insecticides are able to penetrate the waxy cover- regulator (IGR) buprofezin was not decisive;
ing of mealybugs, those containing ethyl alcohol however, the IGR pyriproxyfen and the insecti-
(ethanol), such as some oil-based insecticides, cide onicamid were not directly or indirectly
may allow the material to penetrate through the harmful to the predator C. montrouzieri and para-
waxy covering, killing mealybugs. When apply- sitoid L. dactylopii, indicating that these insecti-
ing high-volume sprays, thorough coverage is cides are compatible with both the natural
imperative, especially when using contact insec- enemies when used together for the control of
ticides, because mealybugs are commonly citrus mealybug in greenhouses and conservato-
located in areas that are not easily accessible, ries (Cloyd and Dickinson 2006).
such as the base of leaf petioles, leaf sheaths, and Systemic insecticides, those that move
leaf undersides. Adding a spreader-sticker to a throughout plant parts, may also be used to
spray solution may be helpful in improving cov- protect plants from mealybug infestations.
erage and penetration. For highly susceptible Applications should be initiated early in the crop-
plants, it may be prudent to routinely spray with ping cycle or before introducing the plants into
either an insecticidal soap or horticultural oil to interiors. Systemic insecticides may be applied as
prevent mealybug populations from reaching either a growing medium drench or granule. It is
outbreak proportions. Also, it is essential to make important to avoid overwatering plants after-
multiple applications when crawlers are present, ward, so that the roots can absorb the active
because eggs will hatch (with the exception of the ingredient. Systemic insecticides, depending on
long-tailed mealybug) over an extended time the type, may be less effective on mealybugs than
period. Insecticides classied as reduced-risk on aphids or whiteies. This may be associated
include insecticidal soaps, horticultural oils, insect with mealybugs not ingesting lethal concentra-
growth regulators, and systemic insecticides. tions of the active ingredient, because they feed
Insecticidal soaps are usually solutions of a within the mesophyll tissues or on plant stems.
synthetic pyrethrin and a plant-safe detergent. As The use of insecticides is the most effective
with oils, the detergent acts as a surfactant and control against the mealybug when applications
spreader for dispersing the pyrethrin evenly, and are timed to coincide with the crawler stage. In
as a mild caustic against the insects. Pyrethrins greenhouse tests, acephate, oxydemeton methyl,
are synthetic analogs of pyrethrum, the natural and kinoprene suppressed populations of both
extract from certain Asteraceae. Caution should mealybug species and prevented crop damage.
be urged with the so-called safe insecticidal Overall reductions of Rhizoecus oridanus
soaps, as some plants are sensitive, particularly (Hambleton) by kinoprene and Ro 103108 were
tender new tissues. comparable to the insecticides acephate and
Horticultural oil, neem oil, and mineral oil are oxamyl (Hamlen 1977). In greenhouse against
effective for mealybug suppression. Horticultural, P.solenenopsis on coleus Solenstemon scutellari-
mineral, or neem oil solutions smother the oides, soil drenching with thiamethoxam, a
insects; so, complete coverage of all sprayed neonicotinoid-based insecticide, provided the
plants is essential. These oils are mixed with highest mealybug control (Willmott 2012).
water and usually a plant-safe detergent for When using pesticides, nymphs are easier to
enhancing the spreading and sticking of the oil. control than mature mealybugs. Insecticides used
The main caution with these oil solutions is that for mealybug control should be rotated to mini-
they should never be applied to plants on hot mize resistance buildup. Insecticides should be
days or in direct sunlight, as to prevent burning of applied using a sprayer that provides complete
tissues. Also, to prevent sun-burning, the chemi- spray coverage of plant. Particularly for mealy-
cal should be applied and allowed to dry in shade. bugs, it is important to totally wet the entire plant,
68 Glasshouse, Greenhouse and Polyhouse Crops 625
including the basal portion. All pesticide labels tion practices and the management program
should always be read and followed. against another pest. The nontarget effects of a
The following insecticides are registered for biological control agent on other benecial or
use against mealybugs in greenhouses: nonpest organisms have to be investigated. The
most suitable host stages may achieve higher
Acephate, Acetamiprid, Azadirachtin, Beauveria rates of parasitism, survival and development,
assiana, Bifenthrin, Buprofezin, Chlorpyrifos, and produce a higher number of progeny consist-
Cyuthrin, Dinotefuran, Fenoxycarb, ing of mainly female parasitoids. The mean tem-
Fenpropathrin, Flonicamid, Imidacloprid, perature of the greenhouse should be maintained
Kinoprene, Parafnic oil, Petroleum oil, at 15 to 30 C for the parasitoids to achieve the
Potassium salts of fatty acids, Spirotetramat, highest developmental rate. Choosing the appro-
Thiamethoxam. priate release time and environmental conditions
can enhance the establishment and effectiveness
of the parasitoid population. The parasitoids can
68.7 Biological control be released as an inundative or seasonal inocula-
tive biological control agent when the mealybug
With some of these chemicals facing phase-out, population level is low. When the mealybug pop-
and with the rising environmental and economic ulation is high, chemical control may be required to
concerns surrounding chemical control tactics, reduce the mealybug population below the dam-
biological control presents a promising alterna- aging level, before the parasitoids can be released.
tive to chemical control for greenhouse ornamen- Insecticides of choice may include insect growth
tal growers. The waxy covering may be the regulators and other compatible chemicals.
reason for the rare occurrence of pathogens and
nematodes as major infesting agents of the
mealybug (Franco et al. 2009). Still, biological 68.7.1 Planococcus citri
control of greenhouse pests through introduction
of natural enemies offers a viable alternative to Biological control agents currently available for
chemical controls. The use of biological control suppression of citrus mealybug populations
agents such as parasitoids and predators has been include the predatory ladybird beetle,
successful in managing mealybugs, primarily cit- Cryptolaemus montrouzieri, commonly referred
rus mealybugs, under specic crop production to as the mealybug destroyer, and the parasit-
systems and interiorscapes. Biological control of oid, Leptomastix dactylopii. The larval stages of
mealybug in greenhouse production relies on the mealybug destroyer resemble mealybug
augmentative releases of parasitoids and preda- adults. L. dactylopii females only attack the third
tors. Biological control agents that are available instar and young adult female life stages. Both
commercially include a variety of tiny parasitic the natural enemies are effective in suppressing
wasps, brown lacewings, green lacewings, and or regulating citrus mealybug populations, and
lady beetles. Some of the commercially available they can be used together under certain systems
mealybug natural enemies are the parasitoids and situations. Doutt (1952) demonstrated that
Anagyrus pseudococci (Girault), Leptomastidea the mealybug P. citri could be successfully con-
abnormis (Girault) and L. dactylopii (Howard) trolled on gardenias by two encyrtid parasites
(all Hymenoptera: Encyrtidae) for P.citri, and the (Leptomastix dactytopii and Leptomasiidea
predator Cryptolaemus montrouzieri (Mulsant) abnormis) and the ladybird Cryptolaemus
(Coleoptera: Coccinellidae) for many mealybug montrouzieri Mulsant. One of the difculties
species (Chong and Oetting 2007). encountered in the use of a predatory insect as an
Biological control in greenhouse ornamental agent of pest control is that the near eradication
production is characterized by the diversity of of the host, in this case mealybugs, is followed
plants and pests. A biological control program by the disappearance of the predator. This
for one pest must be compatible with the produc- necessitates reintroduction of the natural enemy.
626 K.G. Pillai
The parasitic wasps Leptomastix dactylopii and Leptomastidea abnormis. Successful control was
Anagyrus pseudococci are commercially avail- obtained during summer and autumn, but not in
able for the control of citrus mealybugs. winter when the temperature was 13-17 C.
Generalist predators, such as green lacewings Leptomastix dactylopii was more successful in
Chrysoperla spp., and a mealybug predator summer and Leptomastidea abnormis in autumn.
Cryptolaemus montrouzieri are also marketed as Aggregation of adults of Leptomastix dactylopii
biological control agents of mealybugs. occurred at the level of sample areas, but no spa-
Cryptolaemus montrouzieri is highly effective in tial relationship was found between host density
the control of mealybugs in greenhouses. and percentage of parasitism.
Cryptolaemus has also been often used to control Introduction of parasitoids gave improved
the mealybugs in glasshouses. The temperature biological control of P. citri in a large glasshouse
has to be above 20 C in the glasshouse, and the stocked with a variety of ornamental plants in the
mealybug infestation should be great enough to United Kingdom, supplementing that achieved
provide adequate food for the predator (Panis and by the coccinellid predator Cryptolaemus mon-
Brun 1971). Planococcus citri on gardenias and trouzieri. Following the release of parasitoids
Phenacoccus gossypii on chrysanthemum were Leptomastix dactylopii and Leptomastidea
controlled effectively by the release of C. mon- abnormis, there was evidence of mealybug popu-
trouzieri. One adult per plant of gardenia and one lation regulation on guava and coffee bushes
for each two chrysanthemum plants were with reduced and stabilized mealybug numbers
released. C. montrouzieri was recommended to and stable percentage parasitism. The encyrtid
compliment L. dactylopii for the control of orna- Leptomastidea abnormis was responsible for
mentals in the glasshouse. Good control of P. about 90 % of the parasitism observed; the
citri on Clivia and crotons, and reasonable con- remainder was by another encyrtid, Leptomastix
trol on Pelargonii, Saintpaulia, Cattleya, and dactylopii. The combinations of L. dactylopii and
Pilea were observed (Copland et al. 1985). C. other parasitoids (e.g., L. abnormis) and preda-
montrouzieri was used to control P. citri on the tors (e.g., C. montrouzieri) are most effective
crops grown in glasshouses (Lagowska 1995). In against P. citri in greenhouses (Copland et al.
the green net house, Cryptolaemus, when released 1985; Chong and Oetting 2007). Inoculative
at 20 larvae/plant, was found highly effective in release of ve encyrtid parasitoids, Leptomastidea
clearing the mealybugs P. citri on the ornamen- abnormis, Anagyrus pseudococci, L. dactylopii,
tals red ginger, Heliconia, etc. within 2 months of Chrysoplatycerus splendens (Howard), and
its release in India. In Canada, C. montrouzieri Coccidoxinoides perminutus (Timberlake),
was found in greenhouses on P. citri and P. gos- resulted in the rapid suppression of citrus mealy-
sypii (McLeod 1939). P. citri is the major pest of bug, P. citri, on greenhouse citrus. Several para-
ornamental citrus plants in greenhouses. A sites, L. abnormis, A. pseudococci, and L.
predator:prey ratio of 1:15, in most cases, resulted dactylopii, persisted for periods >20 weeks and
in lower populations of P. citri. When compared maintained the host at reduced densities through
with Nephus reunioni (Fursch), C. montrouzieri delayed density-dependent regulation (Summy
caused a signicant reduction in the mealybug et al. 1986; Van Lenteren and Woets 1988).
population. In most cases, signicant differences
in pest reductions were not detected between C.
montrouzieri and methidathion on potted orange 68.7.2 Pseudococcus viburni syn.
plants (Hamid and Michelakis 1994; 1997). P. afnis and P. obscurus
In a commercial greenhouse in Leiden,
Netherlands, biological control of the pseudococ- Good control of Pseudococcus obscurus (Essig)
cid P. citri on Stephanotis plants was carried out on cacti and Clivia were achieved by using C.
with the coccinellid predators Cryptolaemus montrouzieri (Copland et al. 1985). The
montrouzieri and Nephus reunioni, and with the Australian ladybird beetle Cryptolaemus mon-
encyrtid parasitoids Leptomastix dactylopii and trouzieri is used to control the mealybugs in
68 Glasshouse, Greenhouse and Polyhouse Crops 627
Hamlen RA (1977) Laboratory and greenhouse evalua- Panis A, Brun J (1971) Trial of biological control against
tions of insecticides and insect growth regulators for three species of Pseudococcidae (Homoptera,
control of foliar and root infesting mealybugs. J Econ Coccoidea) in greenhouses of ornamental plants.
Entomol 70(2):211214 Revue de Zool Agricole 70:4247
Hayat M (2009) Description of a new species of Aenasius Summy KR, French JV, Hart WG (1986) Citrus mealybug
Walker (Hymenoptera: Encyrtidae), parasitoid of the (Homoptera: Pseudococcidae) on greenhouse citrus:
mealybug, Phenacoccus solenopsis Tinsley Homoptera: density-dependent regulation by an encyrtid parasite
Pseudococcidae ) in India. Biosystematica 3:2126 complex. J Econ Entomol 79(4):891985
Heidari M (1999) Inuence of host-plant physical defenses Van Lenteren JC, Woets JV (1988) Biological and inte-
on the searching behaviour and efcacy of two cocci- grated pest control in greenhouses. Annu Rev Entomol
nellid predators of the obscure mealybug, Pseudococcus 33(1):239269
vibruni (Signoret). Entomologica 33:397402 Waterworth RA, Redak RA, Millar JG (2011) Pheromone-
Lain HM, Parrella MP (2004) Mealybug species baited traps for assessment of seasonal activity and
(Hemiptera: Pseudococcidae) found on ornamental population densities of mealybug species (Hemiptera:
crops in California nursery production. Proc Entomol Pseudococcidae) in nurseries producing ornamental
Soc Wash 106:475477 plants. J Econ Entomol 104(2):555565
Lagowska B (1995) The biological control perspective of Willmott AL (2012) Efcacy of systemic insecticides
scale insects (Homoptera, Coccinea) on ornamental against the citrus mealybug, Planococcus citri, and
plants in glasshouses. Wiadomosci Entomologiczne pesticide mixtures against the western ower thrips,
14:510 Frankliniella occidentalis, in protected environ-
McLeod JH (1939) Biological control of greenhouse ments. Masters thesis. Kansas State University,
insect pests. Rep Entomol Soc Ont 70:6268 Manhattan, KS
Root Mealybugs
69
Maicykutty Mathew and M. Mani
Root mealybugs are several small species of eld crops. The species belonging to genera
mealybugs found below the soil surface, and Geococcus, Rhizoecus, Xenococcus,
feed on root and root hairs in numerous plants. Chorizococcus, Spilococcus, Spinococcus and
They are also called soil mealybugs and subter- Chnaurococcus are known to roots of the
ranean mealybugs. Infestations frequently are plants (Table 69.1).
not detected as the pests occur in the soil, and
populations are quite slow to develop, with
36 months occurring before infestations are
easily visible. Careful examination of infested 69.1 Important Root Mealybug
roots will reveal white, cotton-like masses. Species
These white masses contain both mature
females and eggs. Infected plants become 69.1.1 Gonococcus coffeae
wilted and stunted with foliar yellowing or
chlorosis. They are oval shaped (1/16 to 3/16 Geococcus coffeae Green can be easily be distin-
of an inch long) that look like they have been guished by the pair of stout dorsal spines situated
covered by our. Because they are white or on the head (Green 1933). Geococcus coffeae
light grey in colour, they often resemble small was known to infest sweet potato Ipomoea bata-
grains of rice. These mealybugs have a thin, tas in Tamil Nadu, India (Williams 1985) and
uniform waxy coating and lack the terminal also several other plants such as Theobroma
wax laments typical of their foliar-feeding cacao, Coffea spp., ornamentals, pine apple, and
relatives. Root mealybugs are slow moving, palms (Ben-Dov 1994).
sac-like mealybugs with pronounced crosswise
grooves. They do not have laments surround-
ing their body like many of the foliar feeding 69.1.2 Geococcus citrinus
mealybugs. Root mealybugs pose serious prob-
lem to potted and greenhouse plants and also Geococcus citrinus is a ground mealybug that
lives in the soil and damages the root of citrus in
M. Mathew (*)
Kerala Agricultural University, Trichur, India
e-mail: maycypm@yahoo.co.in
M. Mani
Indian Institute of Horticultural Research,
Bangalore 560089, India
Table 69.1 List of other root mealybugs on different host plants in different countries
Mealybug species Plants Country
Dysmicoccus brevipes (Cockerell) Pigeon pea & groundnut South India
Pineapple Many countries
Dysmicoccus texensis (Tinsley) Coffee Espirito Santo
Cassava Paraguay, Bolivia & Brazil
Dysmicoccus vaccinii sp. n. Blueberries USA
Ferrisia virgata (Ckll.) Parthenium hysterophorus India
Geococcus johorensis Williams Oil palm Johore & Malaya
Geococcus lawrencei Williams Asplenium nidus Solomon Islands
Geococcus oryzae Kuwana Oryza sativa Japan & Ceylon
Phenacoccus salviacus Moghaddam Salvia bracteata Iran
Phenacoccus hordei (Lindeman) Grasses, alfalfa, barley, clover, rye & European countries
wheat
Planococcoides robustus Ezzat & Mango India
McConnell
Planococcus citri (Risso) Coffee Kenya/East Africa
Citrus Crete
Planococcus cryptus Hempel Coffee Brazil
Planococcus cus Signoret Grapevine South Africa
Planococcus fungicola sp. nov. Coffee Kenya
Pseudococcus eriocerei Williams Cacti Argentina
Pseudococcus viburni (Signoret) Plum Chile
Pseudococcus cryptus Hempel Coffee Espirito Santo
Polystomophora arakensis Atraphaxis sp. Iran
Moghaddam
Rhizoecus maasbachi Jansen Segeretia theezans Netherlands
Michelis sp. China
Segeretia sp. England
Rhizoecus amorphophalli Betrem Amorphophallus variabilis Java
Amorphophallus sp. India
Gingiber ofcinale
Diosorea elephantipes
Curcuma domestica
Amorphophallus variabilis Caroline Islands
Colocasia esculenta, Curcuma longa Philippines
and Kaempferia galangal
Rhizoecus theae sp.n. Tea Japan
(continued)
69 Root Mealybugs 631
69.1.17 Chryseococcus arecae per plant was recorded in Gedeo zone while the
lowest infestation of three mealybugs per plant
The golden root mealybug, Chryseococcus are- was recorded in Yem district. Knowledge about
cae Maskell is a native of New Zealand. It was the biology and distribution of this species has
found in Britain and can be witnessed on the paramount importance in devising proper man-
roots of outdoor plants all year round. Golden agement. Enset plants infested with mealybugs
root mealybug is a sap feeding insect that feeds have a retarded growth and dried lateral leaves.
on the roots of a wide variety of plants, although The insects attack all plant age groups but symp-
it has only been found on Meconopsis and toms are more severe on 2 to 4 years old enset
Primula in UK. Mealybug infestations have been plants. Enset root mealybugs are found on roots
noticed on plants lacking vigour. and corms. However, during periods of extreme
drought the mealybugs tend to move towards the
corm when some of the roots drought. The dis-
persal mechanism of enset root mealybugs is
facilitated by movement of infested suckers, farm
implements during cultivation, repeated trans-
planting operations and association with ants.
The population density of the mealybugs was sig-
nicantly (P < 0.05) higher on the roots than the
corms. Enset root mealybugs were found up to a
soil depth of 60 cm and up to 80 cm from the
corm. However, root density as well as mealybug
population numbers decreased with increasing
soil depth. About 99 % of the mealybugs and
Chryseococcus arecae 96 % of the roots were collected within the upper
40 cm soil layer. In addition, about 90 % of the
mealybugs were found within a 60-cm radius
from the plant (Addis et al. 2008, 2010).
69.1.18 The Enset Root Mealybug
Cataenococcus ensete
69.2 Damage
Enset (Ensete ventricosum) was domesticated in
Ethiopia several hundred years ago, and is now There can be several generations of the root
the staple food crop for over 15 million Ethiopians mealybugs throughout the year and numbers can
living in the highlands of southern Ethiopia. The multiply under favourable conditions. With
enset root mealybug Cataenococcus ensete severe infestations, root mealybugs can be found
Williams and Matile-Ferrero is a major pest in on the soil surface at the stem base. It is very dif-
the enset growing regions of southern Ethiopia. cult to detect symptoms of root mealybugs on
Infestation was high in Amaro, Gedeo, Sidama the plant. White, cottony-like masses containing
and Bench districts with 100, 67, 61 and 57 % egg-laying females and/or eggs are normally vis-
incidence respectively. Low mealybug incidence ible on the outside of the root mass when an
was recorded in Gurage, Kembata Tembaro, infested plant is lifted. Slow plant growth and
Hadyia zones and Yem districts. More than 30 % leaf deterioration may be signs of the presence of
of the enset farms were infested with the mealy- the pest. Root-bound or under environmental or
bugs. The highest infestation of 81 mealybugs nutritional stress, the plants are more susceptible
69 Root Mealybugs 635
to attack. Once established in the greenhouse, The adults and nymphs of Geococcus suck sap
root mealybugs may spread as crawlers from from the lateral roots of banana colonizing at the
plant to plant as the water moves out of the drain- junction of laterals with main root resulting in
age holes to nearby plants and in plant debris. It drying up of such roots. Yellowing and narrowing
is mainly potted plants (especially bonsai plants) of leaves, general weakening of the plant, reduc-
that are concerned during import inspections. tion in bunch weight, etc. were the observed
The pot should be removed and roots examined symptoms. Geococcus citrinus occurs seriously
for waxy secretions. In case of heavy infesta- on banana roots in reclaimed paddy elds. G. cof-
tions, crawlers may be observed on the soil sur- feae was also associated with banana grown in
face. The mealybugs may be found particularly in uplands.
the new feeder roots in the upper layer of the soil. The adults and immature stages of Rhizoecus
The resulting damage sties the ability of roots to hibisci feed on plant roots particularly new roots
absorb water and nutrients. The only outward in the upper layer of soil reducing water and
sign of root mealybug feeding may be a decline nutrient uptake by host. Feeding reduces plant
in the health of infested plants. When plants are growth resulting in shrivelling and crinkling.
removed from the pot, the whitish mealybugs Leaves wilt, become pale and turn yellow or
feeding on the roots are then observed. If the grey; alternatively they can become soft, translu-
plant seems to be declining in health because it cent and brown. Flowers may not be produced.
has yellow foliage or slow growth or is stunted Mealy bugs (Planococcus sp., P. citri, P.
for what seems to be no particular reason, then it lilacinus, Dysmicoccus brevipes and Ferrisia vir-
is to be looked for something that could be lurk- gata) were found infesting the roots and basal
ing below feeding on the plants root system. In region of stem of black pepper vines (Piper
case there are mealybugs on bonsai trees, leaves nigrum). Infested plants show slow or poor
may be pale (sometimes greyish) or wilted, growth. Leaves wilt, become pale or turn yellow
despite regular fertilizer and watering. Maybe the or grey. Wax deposit is seen around the roots, on
plant growth has slowed down and/or owering the soil or on the side of the pots. The infestation
has ceased. In severe cases, the leaves may be is generally severe during the post monsoon. The
misshapen. Although they occur throughout the root mealy bug affects the aerial parts of the black
roots, they are most obvious along the edges. pepper vines such as the tender shoots, leaves and
berries (Devasahayam et al. 2010).
Parputo sp. cause appreciable damage to
mulberry directly by sucking the sap and indi-
rectly by making way for some fungal infection,
leading to rotting of the root and ultimately
death of the plants. The infested mulberry plants
show vulnerability to the attack of various fun-
gal pathogens such as Fusarium solani,
Phomopsis mori and Colletotrichum gloeospo-
rioides. Due to this, decaying of bark portion of
root and stem occurs with severe anthracnose
disease. Finally, it results in the death of such
severely affected mulberry plants (Biswas et al.
Mealybugs on the roots
2002).
636 M. Mathew and M. Mani
Roots of banana infested with mealybugs Banana plants infested with root mealybugs
used when replanting. Regular applications in a diluted chlorpyriphos solution (1 pint per
(weekly for several weeks) of insecticide 100 gal) for about 30 s with slight agitation is
watered into the soil are also effective; it is nearly twice as effective as dipping the plant
also possible to immerse the plant pot up to while still in its pot. Imidacloprid, which can
the top of the soil in a bucket of insecticide. be applied only as a drench and incorporated
A promising alternative to chemical treat- with a surfactant or wetting agent to ensure
ments has been found in the use of thorough distribution of solution in the potting
Diatomaceous Earth, a fully inert, non-vola- medium, can also signicantly reduce the
tile substance that has proven effective in number of individuals in an infestation (Hata
eradicating certain insect pests. Strictly speak- et al. 1996).
ing, Diatomaceous Earth is not an insecticide. Moth ball: As a preventative measure, moth
It is made from the skeletal remains of dia- balls (paradichlorobenzene), added to the pot-
toms, a microscopic form of algae. When pro- ting mix, seem to discourage infestation by
cessed into Diatomaceous Earth, these skeletal root mealy bug, and probably discourages
remains form razor-sharp particles which cut other insects. However, the chemicals in the
into the bodies of small insects. While eradi- moth balls can cause damage to plastic plant
cating the insects, Diatomaceous Earth does pots and are best used with clay pots.
not harm African Violets. To treat for soil Traditionally, the only effective treatment for
mealy bugs, repot the African Violet in a soil soil mealybugs (R. amercanus) has been to
that has been mixed with Diatomaceous Earth. spray the soil with acephate (as directed on the
Use about one tablespoon per one litre of soil. label) or with malathion (1 teaspoon of
Pasteurize soil before re-potting. To make soil Malathion 50 per 4 l of lukewarm water).
uninhabitable for future mealy bug infesta- While this treatment does work, it usually
tions, mix about one fourth tablespoon of takes several applications over a period of
Diatomaceous Earth with every litre of soil. days. Moreover, there is usually some risk to
Hot-water dips are as effective as insecticides plants when using any chemical treatment.
against mealybugs. Submerging the potted
palms in water held at 120 F (49 C) until the
internal root ball temperature reached 115 F
(46 C) was 100 % effective in killing root 69.3.2 Field Conditions
mealybugs. Drenching potted palm roots in
hot water at 120 F for 15 min will not only Application of sodium silicate and calcium oxide at
control mealybugs but will also eliminate bur- the time of planting effectively reduced the popula-
rowing nematodes. If an infestation is found tion of banana root mealybug, G. citrinus.
(Rhizoecus hibisci), hot water treatment of Drenching of the chemical insecticides, chlorpyri-
root balls is very effective (Hu et al. 1996). phos at 0.05 % at monthly intervals, reduced the
Chemical control of root mealybugs requires root mealybug population. Among the combina-
saturation of the root ball and potting medium tions, without synthetic insecticides, sodium sili-
to a degree that allows the pesticide to pene- cate alone and its combination with neem seed
trate the pests white, waxy secretion. Dipping kernel extract (NSKE) and Cephalosporium leca-
or drenching with liquid insecticide is more nii Zimm, were effective in reducing the mealybug
effective than applying a granular formula- population at sixth and seventh month of the crop.
tion. Chlorpyriphos, applied twice as a drench Application of chlorpyriphos gave the highest ben-
or dip at 2-week intervals controls coffee root et--cost ratio of 2.46 followed by sodium silicate
mealybug; however, it may take 46 months (2.30) (Smitha and Mathew 2010b). Application of
before the cottony, waxy secretions deterio- neonicotinoids, which include imidacloprid, thia-
rate completely. In the dip method, submerg- methoxam, thiocloprid, by way of soil drench can
ing the plants entire root ball without the pot also be tried against root mealybugs in general.
69 Root Mealybugs 639
fact that the root mealybug females and nymphs Green EE (1933) Notes on some Coccoidae from Surinam,
Dutch-Guiana, with descriptions ofnew species.
are wingless and must crawl to potted plants to
Stylops J Taxon Entomol 2:4958
become established. The procedure is expected to Hamon AB (1982) Rhizoecus arabicus Hambleton, a root
permanently eradicate rather than simply control mealybug in Florida (Homoptera: Coccoidea:
these pests. Pseudococcidae). Entomology Circular, Division of
Plant Industry, Florida Department of Agriculture and
Consumer Services, 238, 2 p
Hara AH, Nino-DuPonte RY, Jacobsen CM (2001) Root
References mealybugs ofquarantine signicance in Hawaii.
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Abraham V, Ajith CB, Priyamol S (2000) Krishiyankanam, of Hawaii, Manoa
6 (4): 56 Nair, M.R.G.K., Visalakshi, A. and Koshy, Hata TY, Hara AH, Hu BKS (1996) Use of a systemic
G. 1980. A new root infesting mealybug of coconut. insecticide granule against root mealybugs, Hawaii.
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distribution of enset root mealybugs on enset. Afr Hu BKS, Hara AH, Hata TY (1996) Hot water as a poten-
Crop Sci J 16(1):6774 tial treatment against root mealybugs, Hawaii, 1995.
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Issue: Bananas, Plantains and ensete II.). Tree and Studies on Paecilomyces lilacinus, an entomopatho-
Forestry Science and. Biotechnology 4(1):3946 gen on Root mealy bug of Banana in Global confer-
Alves VS, Moino Junior A, Santa-Cecilia LVC, Rohde C, ence on banana organized by AIPPUB, ICAR,
da Silva MAT (2009) Revista Brasileira de Bioversity International and NRCB, Trichy, during
Entomologia 53(1):139143 1013 December 2010
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70
P.K. Vinod Kumar, G.V. Manjunath Reddy,
H.G. Seetharama, and M.M. Balakrishnan
70.1 Species Distribution gata Ckll. has also been recorded (Chacko and
Bhat 1976). They attack both robusta and arabica
Among the two commercially cultivated coffee but prefer the former. Planococcus cus and P.
varieties, Coffea arabica L. (arabica coffee) and minor have been recorded on coffee as minor
C. canephora Pierre ex Froehner (robusta cof- pests. The mealybugs, P. citri and P. lilacinus, are
fee), the latter is more prone to attack by mealy- distributed throughout the coffee tracts of India
bugs since this variety is grown in more open and can be noticed quite often during the summer
conditions and at lower elevations. Over 50 spe- months. Planococcus lilacinus is predominantly
cies of scales and mealybugs are reported to found in Kodagu district of Karnataka state,
attack various parts of the coffee tree roots, while P. lilacinus and P. citri are found in equal
branches, leaves, ower clusters and berries proportion in Wayanad district of Kerala state in
where they suck the sap and are of great eco- India (Abdul Rahiman et al. 1995). In Wayanad
nomic importance (Wrigley 1988). Planococcus district of Kerala, the population of P. citri was
kenyae (Le Pelley), popularly known as coffee higher in all the zones compared to P. lilacinus
mealybug, is distributed in Uganda, Tanzania and (Abdul Rahiman and Naik 2009b). For P. lilaci-
Kenya (Bigger 2009). The two most commonly nus, several collateral hosts have been recorded,
encountered mealybugs on coffee in India are which can aid in the survival of the mealybug
Planococcus citri Risso (Coleman and Kannan even if adequate measures are adopted to control
1918; Ayyar 1940) and P. lilacinus Ckll. (Sekhar them on coffee (Bhat and Shamanna 1972)
1964; Bhat and Shamanna 1972). Ferrisia vir- (Table 70.1).
70.3 Seasonal Development and the cock tailed ant, Crematogaster sp.,
actually chase away the bigger predators while
Mealybug population increases if warm and their constant presence over the mealybug colony
humid conditions prevail. Continuous monsoon, is a hindrance for the parasitoids. This is evident
high humidity and low temperatures are detri- in the case of the lepidopteran predator Spalgis
mental to mealybug development. The migration epeus Westwood wherein the aggressiveness of
of mealybugs starts in September/October from the ants and S. epeus population indicated a
the ground to the aerial parts of the coffee plant highly negative relationship. Species belonging
along the main stem. The attack of mealybugs to the genus Crematogaster interfered more with
becomes severe during summer and with the predator activity than the ant O. smaragdina
intermittent showers/irrigation (Anonymous (Vinod Kumar et al. 2008a). About 27 species of
1998). Excessive removal of shade in the robusta ants have been recorded world over in associa-
plantations often leads to are up of mealybugs. tion with different species of homoptera attack-
Planococcus citri on arabica and robusta coffee ing coffee. Thirteen species, namely
is distributed throughout the coffee tracts of Crematogaster sp., Anaplolepis longipes Jerdon,
India, mostly on robusta coffee which is grown at Myrmica brunnea Saunders, Plagiolepis sp.,
lower elevations with lesser shade. Two peaks Paratrechina longicornis Latreille, Camponotus
were in FebruaryMarch and JanuaryMarch; rufogalaucus Jerdon, Anoplolepis gracilipes
there was a positive correlation between maxi- (F. Smith), Tapinoma melanocephalum
mum temperature and adults and nymphs and a (Fabricius), Oecophylla smaragdina (Fabricius),
negative correlation with relative humidity and Acropyga sp., Technomyrmex albipes Smith,
nymphs (Gokuldas Kumar 1987). According to Solenopsis geminata Fabricius, Monomorium
Vinod Kumar et al. (2007), the population of P. sp., have been recorded from coffee tracts of
citri on coffee responded positively to maximum South India (Venkataramaiah and Abdul Rahiman
temperature and had no correlation with mini- 1989).
mum temperature. More than rainfall, relative Of the ant species so far recorded, Plagiolepis
humidity was negatively correlated with the sp. is widespread and seen in almost every estate
mealybug population. The hours of sunshine in the coffee growing regions. Acrophaga sp. is
received had a positive correlation with mealy- recorded from Kodagu district of Karnataka state.
bug population (Vinod Kumar et al. 2007). The presence of ant O. smaragdina along with
mealybugs is not a limiting factor for the estab-
lishment of introduced parasitoid Leptomastix
70.4 Ant Association dactylopii attacking P. citri in the eld. The ant
with Mealybugs species associated with mealybugs recorded from
other coffee growing countries are: Camponotus
Mealybugs produce honeydew, a sweet excretory sp. in Brazil, Lepisiota incise (Forel) in Kenya,
product, to which ants are attracted. Ants provide Myrmelachista ramulorum Wheele, Paratrechina
mealybugs sanitation and protection from natu- jaegerskioeldi (Mayr) in Kenya, Solenopsis
ral enemies. The ants feed on the honeydew and punctaticeps (Mayr) in Kenya (James 1933),
act as clearing agents. The common ants found in Pheidole speculifera (Emery) in Kenya, Lepisiota
association with the mealybugs on coffee in India capensis (Mayr) in Kenya, Monomorium phara-
are Anoplolepis longipes, Oecophylla smarag- onis (Linnaeus) in Kenya, Myrmicaria natalensis
dina and Crematogaster sp. (Venkataramaiah and eumenoides (Gerstaeker) in Kenya, Pseudolasius
Rahiman 1989). Sometimes, ants of the genus gowdei (Wheeler) in Uganda, Pheidole punctu-
Camponotus are also observed. Some of the lata (Mayr) in Kenya and Technomyrmex albipes
aggressive ants like the red ant, O. smaragdina, (F. Smith) in Kenya.
70 Coffee 651
A newspaper may rst be tightly tied on the stem cloprid and thiamethoxam in the liquid form,
and over this paper, grease may be applied. applied to the base of the plant, cause 100 %
Optimum shade maintenance helps in regulating mortality of the coffee root mealybug,
the micro-climate around the coffee plants. Plants Dysmicoccus texensis, independent of the coffee
exposed to sunlight are favourable to mealybug plants age, in a single application (Souza et al.
attack. Since many of the common weeds found 2007).
in the coffee plantations harbour mealy bugs, it is Planococcus kenyae only be controlled by a
best to destroy the weeds regularly. combination of measures. Ant management prac-
tices included banding the coffee plants with
20 cm wide plastic bands covered with a sticky-
70.6.2 Chemical Control substances mixed with insecticide chlorpyrifos.
Removal of suckers that touch the ground is to be
Control of mealybugs on coffee using insecti- done to prevent ants. Spraying on the ant nests in
cides was the choice option before stress was the ground with the insecticides is to be carried
placed on biological control (Rangashetty et al. out to control the ants,
1959). Several trials were conducted using insec- The other management includes the applica-
ticides for achieving affordable control of mealy- tion of oils (such as vegetable oils, neem oil or
bugs (Sekhar and Narayana Rao 1964; Chacko mineral oils) or soapy solutions (12 %) to kill
et al. 1976; Vinod Kumar and Prakasan 1992). mealybugs by suffocation. Spraying cow urine
The insecticides tried were mostly organophos- fermented for 1 day, in a ratio of 1 urine : 4 water
phates. Synthetic pyrethroids did not show any can cause moderate reduction of mealybug popu-
promise against the mealybugs. But most of these lation. Spraying with dimethoate, diazinon, eth-
insecticides were highly toxic to the introduced ion and carbaryl are more toxic (class II,
natural enemies as well as indigenous natural moderately hazardous) (http://www.plantwise.
enemies (Chacko et al. 1979; Stephen et al. 1981; org/FullTextPDF/2013/20137803401.pdf). In
Reddy et al. 1988; Vinod Kumar et al. 2010). In Brazil, with systemic insecticides for the control
the case of severe incidence, quinalphos 20EC at of Dysmicoccus cryptus (Hemp.) (Planococcus
300 ml in 200 L of water plus 200 ml of any wet- cryptus), which attacks the roots of coffee, mor-
ting agent is recommended as hot spot applica- tality was complete and no reinfestation occurred
tion and not as a blanket spray. If the root region for more than 60 days when granules containing
is infested with the mealybug P. lilacinus, a soil 10 % aldicarb had been placed in a furrow (10 cm
drench with dimethoate 30EC at 660 ml in 200 L deep at a radius of 30 cm from the trunk) at the
of water is found to be extremely useful (Vinod rate of 75 g/tree, or when an emulsion spray con-
Kumar and Prakasan 1992). Kerosene, as spray, taining 0.06 % vamidothion was applied to the
can also be used as a milder measure to tackle the foliage at 2 l/tree. Good initial results were also
mealybugs. For spray use 4 L of kerosene in obtained with granules containing disulfoton,
200 L of water along with a wetting agent. The phorate or aphidan [S-((ethylsulnyl) methyl) O,
solution should be mixed thoroughly with the O-bis (1-methylethyl) phosphorodithioate]
wetting agent so that any risk of un-emulsied (Cavalcante 1975).
kerosene falling on the plants is avoided
(Gokuldas Kumar et al. 1989). Plant products,
like neem formulations, have also been tested 70.6.3 Biological Control
against P. citri and some of them have been found
to affect the mealybug population considerably Several indigenous natural enemies on their own
and bring about reduction (Irulandi et al. 2000). are capable of keeping the mealybug population
Iimidacloprid at 0.01 % was known to cause 94 in check (Reddy et al. 1992). This is particularly
% P. lilacinus on coffee after 21 days of spraying true in the case of P. lilacinus, the dipterans
in India (Irulandi et al. 2000). In Brazil, imida- Triommata coccidivora Felt were able to suppress
70 Coffee 653
the mealybug population up to 96 % (Prakasan the predator C. montrouzieri with the perfor-
et al. 1992). mance of the parasitoid L. dactylopii in the eld
In Kenya, the release of C.montrouzieri failed as the predator is not able to discriminate between
to suppress the coffee mealybug Planococcus parasitized and healthy mealy bugs (Prakasan
kenyae. In Celebes, substantial control of and Bhat 1985).
Rastrococcus iceryoides in coffee was obtained The fungus Beauveria bassiana (Bals.-Criv.)
with C. montrouzieri . Control of the mealybug, Vuill. (UEL 114) and the nematode Steinernema
Ferrisia virgata, in coffee plantation of Java was carpocapsae (Weiser) are known to cause high
attempted in 1918 using C. montrouzieri. mortality in short time of adult female mealybugs
Establishment of Cryptolaemus occurred Dysmicoccus texensis (Tinsley) (Andalo et al.
throughout the eastern Java on Planococcus citri 2004). Entomopathogenic nematodes (EPNs)
but with determinable effect on mealybug infes- have potential for biological pest control and
tations which declined. In Dutch East Indies, an have been successfully used in several countries
attempt was made to use C. montrouzieri against in soil and cryptic pests control, as for example
F. virgata on coffee. the coffee root mealybug D. texensis. Greenhouse
In India, severe infestations of mealybugs results demonstrate that aqueous suspension
(Planococcus spp.) occurred in many estates in (JPM3) was more efcient with 70 % control
South Wayanad, Kerala. At Shevaroy hills, adults efciency. In eld experiments, treatments with
and grubs of C. montrouzieri were seen on San aqueous suspensions of insecticide Actara 250
Ramon hybrid coffee where mealybug infestation WG (thiamethoxam), used for comparison, and
was virtually cleaned up (Chacko 1979). A JPM3 were the only ones statistically different
release rate of ve beetles per mealybug infested from control (Alves et al. 2009). Heterorhabditis
Robusta coffee, three beetles per Arabica coffee bacteriophora Poinar strain HC1 was known to
and two beetles per San-ram Coffee plants has cause 100 % mortality in the inoculated the cof-
been recommended to control the coffee mealy- fee mealybug complex (Rodriguez et al. 1997).
bugs in India (Singh 1978). The drawback is that
C. montrouzieri becomes active when the mealy-
bug population reaches high levels by which time References
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loss (Chacko 1982). Leptomastix dactylopii
coffee ecosystem of Wayanad district in Kerala.
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citri, was introduced into India during 1983 from Abdul Rahiman P, Naik PR (2009b) Distribution of coffee
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Biological Control, now the National Bureau of
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has established within two months of release. root mealybug Dysmicoccus texensis (Tinsley)
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(Rhabditida, Heterorhabditidae). [Portuguese].
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