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The tea mosquito bug, Helopeltis theivora Waterhouse (Heteroptera: Miridae):

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The tea mosquito bug, Helopeltis theivora

Waterhouse (Heteroptera: Miridae): its status,
biology, ecology and management in tea plantations
a a b
Somnath Roy , Narayanannair Muraleedharan , Ananda Mukhapadhyay & Gautam
a
Handique
a
Department of Entomology, Tea Research Association, Tocklai Tea Research Institute,
Jorhat, India
b
Entomology Research Unit, Department of Zoology, University of North Bengal, Siliguri,
Darjeeling, India
Published online: 24 Apr 2015.

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 International Journal of Pest Management, 2015
http://dx.doi.org/10.1080/09670874.2015.1030002

The tea mosquito bug, Helopeltis theivora Waterhouse (Heteroptera: Miridae): its status,
biology, ecology and management in tea plantations
Somnath Roya*, Narayanannair Muraleedharana, Ananda Mukhapadhyayb and Gautam Handiquea
a
Department of Entomology, Tea Research Association, Tocklai Tea Research Institute, Jorhat, India; bEntomology Research Unit,
Department of Zoology, University of North Bengal, Siliguri, Darjeeling, India

(Received 3 December 2014; final version received 12 March 2015)

Helopeltis theivora Waterhouse (Heteroptera: Miridae) or the tea mosquito bug (TMB) is a major sucking pest of tea
(Camellia sinensis L.) in most tea-producing countries. The nymphs and adults of the TMB suck the sap from tender
leaves, buds and young shoots, which results in heavy crop losses. The damage to tea plants caused by the TMB is not
limited to the sucking of plant materials and extra-oral digestion by salivation. Damage is also caused by the insertion of
eggs into plant tissues during oviposition. More than a dozen alternate host plants and only a few natural enemies of the
TMB have been recorded. For control of this pest, synthetic pesticides are widely used. There are many reports regarding
the excessive application of insecticides that has resulted in the development of high tolerance and even resistance in this
pest to certain insecticides. This review collates the most important works conducted on the bio-ecology of the TMB and
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includes a discussion of the problems encountered in the management of this pest. The scope of future studies and the
plans for better management of this serious sucking pest of the tea plant are also discussed.
Keywords: Helopeltis theivora; Miridae; tea; bionomics; natural enemies; IPM

Introduction has been no comprehensive review on this major insect

Tea, Camellia sinensis (L.) O. Kuntze, is an intensively
managed perennial monoculture crop cultivated on large-
scale and small-scale plantations. At present, tea is grown
in more than 50 countries around the world and is distrib-
uted from Georgia at 43 N latitude to Nelson (South
Island) in New Zealand at 42 S latitude. Tea is found
from sea level to 2300 m. The prime producers of tea are
China, India, Sri Lanka and Kenya. Vietnam, Turkey,
Indonesia, Argentina, Japan, Bangladesh, Malawi,
Uganda and Tanzania produce the rest of the world tea
crop. The tea crop has unique characteristics that influence
pest ecology in special ways (Calnaido 1973). Tea is an
evergreen and a perennial (over 100 years) crop that is
grown as plantations (Banerjee 1983), with genetically
diverse cultivars and the interplanting of shade trees, par-
ticularly in South east Asia (Deka et al. 2006). Tea planta-
tions roughly resemble a “single species forest” (Cranham
1966a, 1966b), and insect and mite species coexist by
minimizing competition through intra-tree distributions or
well-defined stratification and/or ecological niche forma-
tion (Banerjee 1979, 1983). Worldwide, there are several
species of mirids that are associated with tea. Among the
mirid bugs associated with tea, Helopeltis theivora Water-
house (Heteroptera: Miridae) or the tea mosquito bug
(TMB) is the predominant pest (Cranham 1966b; Hazar-
ika et al. 2009, Saha and Mukhapadhyay 2013). Scientists
from many research institutes around the world are
involved in studying the biology and ecology of this pest
to develop suitable techniques for its management. There
pest of tea. Therefore, the objective of this review is to
consolidate the existing information on the taxonomy,
bio-ecology, pest status, pesticide tolerance, and some
aspects of the integrated pest management (IPM) pro-
grams of this pest. Moreover, we contemplate and discuss
future advances in research that are necessary to improve
the management of the TMB on tea crop.
Taxonomy
The genus Helopeltis is readily distinguished from other
members by the large, spine like process on the scutellum
and by the structure of the male and female genitalia, par-
ticularly of the genital chamber of the female. The TMB
is distinguished from its congeners by a number of charac-
ters, i.e., the base of the narrow, pale antennal segment I,
and the long, erect setae at the distal third of antennal seg-
ment II and much of segment III of the male (Stonedahl
1991). The head has a broad, pale lateral stripe, and the
anterior half of the pronotum and the anterolateral mar-
gins of the disc are pale. The posterior margin of the disc
is infuscated. The lateral edges of abdominal sterna I-IV
are uniformly pale (Stonedahl 1991). Variations in color
in the populations of TMB collected from South India,
Assam and the Dooars were reported by Mann (1907) and
Bora and Gurusubramanian (2007). Rebijith et al. (2012)
successfully conducted the molecular identification of the
TMB by using the mitochondrial cytochrome oxidase-I
(COX-1) DNA sequence. Molecular diagnostics as the

*Corresponding author. Email: director@tocklai.net

Ó 2015 Taylor & Francis

 2 S. Roy et al.
basis for species identification is helpful because the
approach is not limited by developmental stage or gender
(von Dohlen et al. 2006). According to the latest report by
Schuh (2002 2013) [Online Systematic Catalog of Plant
Bugs (Insecta: Heteroptera: Miridae). http://research.
amnh.org/pbi/catalog/], H. febriculosa Bergroth, H. oryx
Distant, H. theobromae Miller, H. theivora theobromae
Miller and Afropeltis theivora are all synonymous with H.
theivora Waterhouse (Stonedahl 1991). This species also
has several common names, viz., mosquito tea bug, mos-
quito bug, tea bug and tea mosquito, and in the present
review, we refer to it as TMB.
Geographical distribution
The genus Helopeltis has a distribution in both northern
and southern hemispheres that include Southeast Asia,
Africa and northern Australia (Stonedahl 1991; CAB
1992). The TMB was first recorded on tea in Java in 1847
(Rao 1970) and on tea in India in 1968 in the Cachar
region (Watt and Mann 1903). In addition to occurring in
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India (Sudhakaran and Muraleedharan 2006; Hazarika et
al. 2009; Saha and Mukhopadhyay 2013), the TMB is
known to occur in most of the tea-producing countries,
i.e., Bangladesh (Ahmed et al. 1992, 2005, 2013), Sri
Lanka (Cranham 1966a, 1966b), Java and Sumatra
(Koningsberger 1908; Leefmans 1916), Laos (Du Pasquier
1932), Vietnam (Du Pasquier 1932), West Malaysia (Lever
1949; Corbett 1930), Taiwan (Hsiao 1983), Papua New
Guinea (Smith et al. 1985), Uganda (Hargreaves 1936),
Kenya (Rattan 1992) and China (Schuh 1995). On Hainan
island of China, the TMB is recorded as a serious pest of
tea and cashew plantations (Yong-ming and Qi-An 1985).
Host range
Tea is the most preferred and principal host of the TMB
(Mukhopadhyay and Roy 2009; Roy et al. 2009h; Saha et
al. 2012a). In addition to tea, it feeds on a wide range of
economic plants including cashew, Acacia, cocoa, cam-
phor and pepper to which it causes considerable damage.
The TMB also subsists on a number of noncrop host
plants, which support their populations when the major
hosts are scarce or have been treated with pesticides. This
behavior enables them to breed throughout the year or to
survive in adverse seasons until the major host is available
in abundance ((Mukhopadhyay and Roy 2009; Roy et al.
2009h). The large number of crop and noncrop host plants
that grow in tea fields or in close vicinity are listed in
Table 1. The complete life cycle of the TMB on alterna-
tive weed hosts such as Maesa indica (Sudhakaran and
Selvasundaram 2000; Sudhakaran and Muraleedharan
2006), Mikania micrantha (Mukhopadhyay and Roy
2009; Roy et al. 2009h; Saha et al. 2012a), Duranta
repens (Gogoi et al. 2012), Psidium guajava (Kalita et al.
2000), Gardenia jasminoides (Kalita et al. 2000), Eugenia
jambolana (Kalita et al. 2000), Melastoma malabathricum
(Kalita et al. 2000) and Chromolaena odorata (Srikumar
and Bhat 2013) have also been reported. The life history
traits of the TMB have changed much with its almost
obligatory dependence on tea. No alternate hosts have
been found to support better performance of the TMB
than tea. The fecundity, rate of growth and survival on tea
were much higher compared with many other hosts, which
proved conclusively that tea is the best of the known hosts
for TMB. Saha et al. (2012a) interpreted the differential
activity of the three principal xenobiotic detoxifying
enzymes [the general esterases (GEs), the glutathione S-
transferases (GSTs) and the cytochrome P450 monooxy-
genases (CYPs)] as causing host-based variation fitness
traits. The activities of these enzymes in the TMB were
significantly enhanced when the insect fed on the alternate
hosts M. micrantha and P. guajava compared with tea,
demonstrating again the suitability of tea as a host.
Nature of damage
The TMB causes serious damage to tea plantations both in
the quantity and in the quality of the tea. The nymphs and
adults of the TMB suck cell sap from tender stems, young
leaves and buds, which results in the formation of reddish
brown circular feeding punctures. The tea buds, shoots
and tender leaves, which are the actual crop of tea,
become curled, dried and black with many sucking stains
and provide no yield. The damaged buds cannot be
plucked (harvested), which affects the next flush of
shoots. The most seriously affected tea plants have a
darker green color and are stunted (Das 1965; Hazarika
2009). Additionally, oviposition causes the stems to
develop cracks and causes over-callusing, which also
results in stunted growth and the dieback of stems (Das
1965; Roy 2008; Sudhakaran 2000). An infestation by
TMBs often starts from a small area in the tea field and
then spreads to patches of neighboring plants. This gives
the tea field an appearance of uneven development (Das
1965). In summary, two types of damage are caused by
the TMB, direct loss of the harvestable shoots and acute
debilitation of the bushes leading to dieback, which delays
flushing and results in poor yields (Rao 1970).
Assessment of crop loss and damage potential in tea
TMB is one of the worst enemies of tea and has caused
losses up to 55% of the crop (Rattan 1992) loss in Africa
and 11% 100% loss in Asia (Muraleedharan 1992a,
1992b). According to Peal (1873), the TMB was destined
to become a major pest of Indian tea plantations causing
enormous damage to the crop within 20 years of its dis-
covery. Before the 1950s, crop losses of 100% were com-
mon because of TMB attack (Rao 1970; Muraleedharan
1987, 1992a), but the use of DDT provided satisfactory
control. However, once the use of this insecticide was cur-
tailed or discontinued, the TMB regained its pest status on
tea (Banerjee 1983; Das 1984). Prasad (1992) and Barbora
and Singh (1994) reported that the usual crop loss due to
TMB infestation was 25% 50%. Earlier studies indicated
that crop loss in a field caused by the TMB could be total
if the attack was severe (Rao and Murthy 1976). In India,
 International Journal of Pest Management 3

Table 1. List of hosts of Helopeltis theivora present in and around tea plantations.

Host plant species Family Reference

Acacia mangium Willd. Fabaceae Thu et al. (2010)

Acalypha indica L. Euphorbiaceae Das (1965)
Anacardium occidentale L. Anacardiaceae Ambika and Abraham (1983),
Sundararaju (1993) and Das (1984)
Annona reticulata L. Annonaceae Kalita et al. (2000)
Anthocephalus cadamba (Roxb.) J. Bosser Rubiaceae Saha and Mukhopadhyay (2013)
Azadirachta indica A. Juss. Meliaceae Sundararaju and Babu (1999)
Bidens pilosa L. Asteraceae Saha and Mukhopadhyay (2013)
Bixa orellana L. Bixaceae Das (1984)
Camellia japonica L. Theaceae Sudhakaran and Muraleedharan (2006)
Camellia sinensis (L.) Kuntze Theaceae Das (1965)
Cannabis sativa L. Cannabaceae Gogoi et al. (2012)
Capsicum spp. Solanaceae Anonymous (2007)
Ceiba pentandra (L.) Gaertn. Malvaceae Das (1984)
Chromolaena odorata (L.) R.M. King & H. Rob Asteraceae Srikumar and Bhat (2013)
Cinchona officinalis L. Rubiaceae Das (1984)
Cinchona sp. Rubiaceae Das (1984)
Cinnamomum camphora (L.) J. Presl. Lauraceae Das (1984)
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Clidemia hirta (L.) Don Melastomataceae Ragesh (2013)

Duranta repens L. Verbenaceae Gogoi et al. (2012)
Ehretia acuminata R. Brown Boraginaceae (Ehretiaceae) Barbora and Singh (1994)
Eugenia jambolana (Syzygium cumini) Myrtaceae Kalita et al. (2000)
Eurya acuminata D.C. Theaceae Das (1965)
Ficus benjamina L. Moraceae Gogoi et al. (2012)
Ficus hispida L.f. Moraceae Saha and Mukhopadhyay (2013)
Gardenia jasminoides Ellis Rubiaceae Kalita et al. (2000)
Ixora coccinea Rubiaceae Gogoi et al. (2012)
Jasminum sandens Vah Oleaceae Das (1965)
Maesa indica (Roxb.) DC. Myrsinaceae Sudhakaran and Muraleedharan (2006)
Maesa ramentacae Wallich Myrsinaceae Das (1965)
Melastoma malabathricum L. Melastomataceae Das (1965)
Mikania micrantha H.B. & K Asteraceae Somchowdhury et al. (1993)
Morus alba L. Moraceae Barbora and Singh (1994)
Murraya koenigii Rutaceae Gogoi et al. (2012)
Oxalis acetosella L. Oxalidaceae Barbora and Singh (1994)
Persea bombycina Kost Lauraceae Gogoi et al. (2012)
Persicaria ( D Polygonum) chinensis (L.) Nakai Polygonaceae Saha and Mukhopadhyay (2013)
Phlogacanthus pubinervius T. Anderson Acanthaceae Somchowdhury et al. (1993)
Phlogacanthus thyrsiflorus (Roxb.) Nees Acanthaceae Gogoi et al. (2012)
Piper hamiltonii C. DC. Piperaceae Gogoi et al. (2012)
Piper nigrum L. Piperaceae Das (1984)
Premna latifolia Roxb. Verbenaceae Barbora and Singh (1994)
Psidium guajava L. Myrtaceae Saha et al. (2012)
Pteridium aquilinum (L.) Kuhn Dennstaedtiaceae Saha and Mukhopadhyay (2013)
Sida cordifolia L. Malvaceae Gogoi et al. (2012)
Smilax herbacea L. Smilacaceae Somchowdhury et al. (1993)
Theobroma cacao L. Malvaceae Anonymous (2007)

the TMB has now attained national importance as a pest,
and it has been estimated that 80% of the tea growing area
is affected, with crop losses of 10% 50% (Bora and
Gurusubramanian 2007; Roy et al. 2008e; Roy and
Gurusubramanian 2013). Ahmed (1996) studied in detail
the loss in tea crop due to TMB infestation in Bangladesh.
He reported an average crop loss of 150 kg tea per ha
(Ahmed 1996). At present in Bangladesh, 10% 15% of
tea crop is lost annually due to the TMB, with losses up to
100% in some cases (Ahmed et al. 2011).
The economic threshold level (ETL) for the TMB in
tea was reported to be a 5% infestation in south Indian tea
 4 S. Roy et al.
plantations (Muraleedharan and Selvasundaram 2002). In
a North Bengal Dooars plantation, Somchoudhury et al.
(1993) reported that the presence of one pair of TMBs in a
group of 10 bushes might cause economic damage to the
crop within 14 days. Sarmah et al. (2011) estimated the
EIL and ETL for a TMB infestation on tea in Assam to
be 3.75% and 2.81% shoot infestation, respectively. The
ETL of the TMB in Bangladesh was a 5% infestation
(Mamun and Ahmed 2011). The ETLs are, however, sub-
ject to change with crop phenology, weather conditions,
cost of control, and market prices of tea, which vary from
region to region.
Life history and biology
Extensive studies conducted on the biology of the TMB
(Das 1965; Gope and Handique 1991; Mann 1902; Sudha-
karan and Muraleedharan 2006; Roy et al. 2009b) have
provided an account of the life history performance of this
pest in India. An individual typically completes its life
cycle on a single bush. The duration of the life cycle var-
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ied with season depending on climatic conditions, from
approximately two weeks in June and July to 5 8 weeks
in colder weather (Das 1984). Details on the biology of
the TMB were reported by Gope and Handique (1991) on
tea from Assam and by Roy et al. (2009e) on tea from the
Dooars of India. The bio-ecology of the TMB in South
India was studied by Sudhakaran and Muraleedharan
(2006). The TMB, in general, has overlapping genera-
tions. In northeast India, the life table of the TMB was
developed by Kalita et al. (1996). In this study, the popu-
lation increased with the intrinsic rate (rm) of 0.152,
whereas the finite rate of increase (λ) was 1.164 per
female per day. The net reproductive rate (Ro) was 28.58,
and the period required for the completion of a generation
was 22.05 days. The rate of population increase was
2.9 times per week, and the population was approximately
93.10% immature stages on reaching a stable age
distribution.
Adults
Das (1957) observed sexual dimorphism in the TMB. The
male was smaller than the female and was slim with a
black pronotal area and a bluish abdomen. The female
was bigger and had a distinct orange pronotum in the tho-
racic region. The abdomen of the female bug was a dirty
whitish green or green (Figure 1).
Color variability in the populations collected from
Vietnam, South India, Assam and the Dooars, with special
reference to the head and pronotum, was reported by
Stonedahal (1991), Mann (1907), Bora and Gurusubrama-
nian (2007), Sarmah and Bandyopadhyay (2009) and Roy
et al. (2009f). Mann (1907) found a strong correlation
between the color variation in the TMBs and the season,
with much darker males in the summer/autumn (July
October) populations than in the winter/spring brood
(November June). The females, however, were reported
to have the reverse coloration. Bora and Gurusubramanian
(2007), Sarmah and Bandyopadhyay (2009) and Roy et al.
(2009f) observed that color variation was present both in
males and females in the same season. Sarmah and Ban-
dyopadhyay (2009) detected genetic polymorphisms using
random amplified polymorphic DNA-polymerase chain
reaction (RAPD-PCR) among the color variants and
showed that the TMB population consisted of discontinu-
ous phenotypes among individuals within a freely inter-
breeding population. The color variants within the TMB
population were different forms of the same species (Sar-
mah and Bandyopadhyay 2009).
The females of the TMB mated four days after emer-
gence whereas males were ready for mating within two
days (Sudhakaran and Muraleedharan 2006). It was the
male that located a female for mating, and copulation
occurred with the abdominal tips of the two sexes fixed
and the heads facing opposite directions. The time of
copulation ranged from 60 to 210 minutes (Sudhakaran
and Muraleedharan 2006; Roy et al. 2009e). Both polyg-
amy and polyandry were common among the TMB, and
some females mated 6 8 times during adulthood (Jeevar-
atnam and Rajapaskse 1981). Peaks in the mating activity
occurred between noon and evening. Gope and Handique
(1991) reported that, under the agro-climatic conditions of
Assam, the preoviposition period for the TMB gradually
decreased with an increase in ambient temperature. How-
ever, a recent study revealed that the preoviposition period
in TMBs was more or less 4 days without much variation
in the different months of the year in the Dooars region
(Roy et al. 2009e).
The young tea shoots are mostly preferred for oviposi-
tion, with occasional laying of eggs in the petioles and
midribs of leaves (Mann 1902; Das 1984). The average
longevity of females was 48 days, whereas males lived for
only 28 days (Sudhakaran and Muraleedharan 2006).
Eggs
The females make a slit with the ovipositor and deposit an
egg separately, which remains completely embedded in
the plant tissue. The dull, white egg is oblong-shaped,
approximately 0.8 1.0 mm long and has two unequal
chorionic processes (Figure 2). The eggs are laid in the
shoot, and the chorionic processes project outside with the
longer process above the shorter one (Gope and Handique
1991), usually with the operculum and respiratory horns
exposed (Cheramgoi 2010). Common sites for oviposition
are the succulent stems of growing shoots and buds at the
leaf axils, and occasionally, the midrib of the leaf,
petioles, broken ends of the plucked shoots or other soft
parts of the bush (Das 1965). Roy et al. (2011) reported
that the most common and preferred site of oviposition
for the TMB was the internode between the first leaf and
second leaf of the shoots, which harbored 28.5% of the
total number of eggs laid, followed by the internode
between the second leaf and third leaf, the stem between
the third leaf and fourth leaf, the lower side of the midrib
and petiole of the fourth leaf, the lower side of the
midrib and petiole of the third leaf, the lower side of the

Figure 1. (Color online) Common morphs of TMB infesting tea plantation in India.
Downloaded by [Somnath Roy] at 20:10 24 April 2015
midrib and petiole of the second leaf, and axillary buds.
The remaining parts of the tea shoot had much fewer
eggs. Therefore, the estimation was that 65.7% of the
eggs were laid in the pluckable portions of the tea, and the
rest of the eggs, 34.3%, were inserted into nonpluckable
regions, i.e., the broken ends of plucked shoots (Roy and
Mokhopadhyay 2011). In contrast, Sudhakaran and Mura-
leedharan (2006) reported that the most preferred site for
oviposition was the broken end (stub) of the stems after
plucking the shoots. Das (1965) reported that a single
female can lay a maximum of 220 eggs in 36 days, with
4 10 eggs per day. The maximum number of eggs laid by
a female ranged from 136 to 180 during September, and
the minimum number of eggs laid was 73 and 74 during
January and February, respectively. Normally, females
laid an average of 6.7 eggs per day (Sudhakaran and Mur-
aleedharan 2006; Roy et al. 2009e). The incubation period
varied from 5 days in August to 16 days in December. In
general, the incubation period is less than a week
Figure 2. (Color online) Eggs of TMB.
International Journal of Pest Management 5
during the months of June, July, August and September
and two weeks or more during November, December, Jan-
uary and February (Gope and Handique 1991; Roy et al.
2009e).
Nymphs
There are five nymphal instars (Figure 3). The young
nymphs emerge by pushing aside the cap of the egg, and
the egg shell remains embedded in the plant tissue (Das
1965). The nymphal period varies from 8.4 to 16.2 days.
The freshly hatched nymph is a dirty yellow with bright
pink antennae and eyes. Soon after hatching, the nymph
begins to feed on the young tender leaves and buds. The
first and second instar nymphs measure approximately 1.5
and 2 mm, respectively, and are greenish yellow. How-
ever, they become green in the later instars. In the first
instar, the eyes are pink, the antennae are longer than the
body, and the labium extends to half the abdomen. In the
 6 S. Roy et al.
Figure 3. (Color online) Different nymphal instars of TMB.
second instar, the drumstick-like process appears. The
third instar is approximately 3 mm long and has a reddish
green body with newly formed wing buds. The fourth
instar is 4 mm long with a greenish yellow body, and the
wing pads become bigger and darker than in the third
instar. The fifth instar is reddish green with a green abdo-
men and is approximately 5 mm long (Das 1965), and the
scutellar process and the wing pads are well developed,
and the sexes can be identified. The males have blunt
abdominal tips, whereas the females have a groove for a
future ovipositor. From May to October, nymphal devel-
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opment is completed within a short time, whereas it is lon-
ger in the month of January (Roy et al. 2009e). Gope and
Handique (1991) reported that the longest nymphal devel-
opmental period was 29.6 days and, in exceptional cases,
may extend up to 39 days (Das 1965).
Seasonal incidence and abundance
Climatic factors such as rainfall and humidity are known
to greatly influence the population dynamics of Helopeltis
spp. (Swaine 1959; Pillai et al. 1979; Geisberger 1983;
Muhamad and Chung 1993; Karmawati et al. 1999). Helo-
peltis was more active early or late in the day compared
with the midday (Watson et al. 1975; Zeiss and Braber
2001). In north east India, Das (1957) reported that the
TMB attack began in tea in May, June and July and often
extended to September when there were more rainy days.
Muraleedharan (1992a, 1992b) provided detailed informa-
tion on the seasonal prevalence of the TMB in southern
India. In southern India, the incidence of the TMB was
high during July to December and low during January to
June. Two peak populations, i.e., June July and
August September, were observed in a year, which coin-
cided with the tea flushing seasons. Preoviposition, incu-
bation and nymphal periods showed significant negative
correlations with morning humidity. Fecundity, hatching
percentage, adult eclosion, reproductive success, total
growth index and sex ratio were positively correlated with
temperature. The seasonal biology of the TMB in north-
east India was studied by Kalita and Singh (1999). The
optimal season for the development of the pest was
August October, followed by February April and
May July. Ahmed et al. (1992) developed a population
model for the TMB in Bangladesh. Chakraborty and
Chakraborty (2001) stated that the attack of tea leaves by
the TMB was positively correlated to temperature and
rainfall, and partially to humidity, for 12 tea varieties
during a two-year period. According to Zeiss and Braber
(2001) and Damiri (2002), the TMB causes severe dam-
age in the humid months of the rainy season in Indonesia
and Vietnam. In a recent study, Roy et al. (2009) reported
that the population of TMBs was abundant throughout the
year in a sub-Himalayan Dooars tea plantation.
According to Das (1965), the following factors are
known to influence the abundance of the TMB: (1) the
nature of pruning and skiffing (removal of their branches
at a specific height from the top of bush), (2) the time of
pruning and skiffing, (3) the degree of pruning, (4) the
height of plucking, (5) defoliation, (6) the shade status,
(7) abandoned tea and seed trees (alternate hosts), (8) the
types or “Jats” (natural variety classified as Chinese,
Assam and Cambod) of tea, and (9) the type of manure. In
conclusion, prolonged wet or humid weather or intermit-
tent rains with warm conditions during the flushing season
normally increase the incidence of the TMB.
Behavior and adaptation
On a cloudy day, the TMBs may be found on the top
branches of the bushes at any time of the day. When dis-
turbed, the nymphs and even the adults quickly move
down the stems or fall to the ground. The adults rarely
migrate unless disturbed. Although adults of TMB are the-
oretically capable of flight for up to 4 km (Cheramgoi
2010), the spread of the bug is mainly affected by winds
carrying them long distances (Das 1965).
Feeding behavior
Both the nymphs and adults of TMBs use their mouthparts
to pierce the young shoots and suck the sap of the young
leaves, buds and tender stems. As the TMBs feed, toxic
saliva is injected that causes the breakdown of tissues sur-
rounding the puncture. Within 2—3 h of feeding, a circu-
lar spot forms around the puncture, and within 24 h, the
area becomes translucent and light brownish and eventu-
ally develops dark brown, sunken spots and dries up. The
affected leaves become deformed and can curl up
(Figure 4). Studies conducted on the nocturnal and diurnal
feeding activities of the TMB revealed that the number of
feeding punctures/shoot was much greater during the
night than during the day (Sana and Haq 1974). A single
fully grown fifth instar nymph was the most voracious
feeder among the life stages and produced the most and
the largest feeding lesions (Bhuyan and Bhattacharyya

Figure 4. Feeding damage by adult TMB on tea: (A) puncture marks after 2 h of feeding, (B) magnified image after 3 h of feeding, (C)
highly infested tea leaf after 24 h of feeding, and (D) badly affected deformed and curled-up leaf after 5 days of feeding.
2006). However, an adult can make 150 feeding spots in a
day (Hainsworth 1952), and a single female can produce
lesions over an area of 412 mm2/day (Kalita et al. 1995).
Severe attacks of the TMB adversely affect the quality
parameters of tea such as the leaf appearance, briskness
Downloaded by [Somnath Roy] at 20:10 24 April 2015
and flavor.
The feeding mechanisms of the TMB were studied by
Cohen-Stuart (1922) who showed that the proboscis pene-
trated the epidermal tissue and caused the collapse of
parenchymal tissue. The stylets transiently punctured epi-
dermal, mesophyll, and parenchyma cells, and this
mechanical damage may influence plant responses to the
TMB infestation (Tjallingii and Esch 1993). Leach (1935)
reported that primary lesions can be recognized before the
insect stopped feeding, which indicated an extremely
active feeding process and phytotoxic salivary secretions.
This rapid feeding was associated with the extensive dam-
age inflicted by small populations of H. theivora. A care-
ful examination of the puncture spots revealed that an
indentation and a reduction in the thickness of the lamina
were associated features of an infestation. Such a progres-
sion may be attributed to hypersensitive reactions
(Fernandes 1990) and subsequent necrosis of adjoining
tissues (Klingler et al. 2005; Gao et al. 2008) following
herbivory.
It is apparent that infestation by the TMB induces
morphological changes in the tea plant, which is affected
by both the mechanical act of infestation and the resulting
hypersensitive reactions that are induced as part of the
direct defense strategy of plants (Bandyopadhyay 2012).
The reactions in the plant are highly pronounced and their
manifestations appear after 30 minutes of feeding. The
ultrastructural changes in the affected cells start with the
collapse of the vacuole and progress to degenerative
changes in chloroplasts and damage to the cell membrane
and cell wall (Ahmed et al. 2013).
The biochemical response of tea to attack by the insect
was determined by Chakraborty and Chakraborty (2001)
who paid special reference to oxidative enzymes and fla-
vonoid and flavor components. According to their obser-
vations, the TMB attack led to an increase in the activities
of the oxidative enzymes peroxidase, ascorbate peroxidase
and polyphenol oxidase. The activity of phenylalanine
International Journal of Pest Management 7
ammonia-lyase generally decreased as a result of this pest
attack. The decrease in the quality constituents of tea
made from infested shoots was mainly because of the
reduction in the important biochemical constituents, such
as total phenol content, and the prior oxidation of the
infested parts. The presence of both hydrolyzing and oxi-
doreductase enzymes in the salivary and midgut homoge-
nates may be related to extra-oral digestion and defense,
leading to tissue necrosis and phytotoxic effects in the tea
leaves (Sarker and Mukhopadhyay 2006). Saikia et al.
(2011) stated that a H. theivora infestation significantly
regulated the content of hormones (auxin and gibberellic
acid), including a stress hormone (abscisic acid). The
damage to the tea plant caused by the TMB is not only
due to sucking plant sap but is also caused by oviposition.
A mated female TMB inserts eggs singly inside the tissues
of the succulent stem by splitting it open with the oviposi-
tor. This results in cracks and over-callusing that lead to
blockages of the vascular bundles, which affects the phys-
iology and causes stunted growth and sometimes dieback
of the stems (Sundararaju 1999; Rahman et al. 2006,
2007).
The rates of feeding and the selection of sites for suck-
ing by nymphs and adults of the TMB were studied in the
laboratory by Kalita et al. (1995) and Rahman et al.
(2007) who reported that the rate of feeding by the female
was higher compared with males. Recently, Roy et al.
(2009d) found a number of lesions in the form of “fluid-
soaked feeding spots” produced by nymphs and adults on
the upper surface of tea leaves. The second leaf was the
most preferred site for the third and fourth instars and the
adults. However, the first and second instars preferred the
first leaf. The assessment of damage potential of the TMB
nymphs and adults in the field under caged conditions was
reported by Dhar et al. (2001) who found that the percent
of shoot damage caused by the adults is higher than that
caused by the nymphs. Because of the oviposition instinct
of adults, they do not attack the same shoot where the
eggs were laid (Dhar et al. 2001) and hence become more
dispersed in the plantation. However, with the nymphs
devoid of wings, there was limited tendency to disperse
and they frequently and repeatedly feed on the same
shoots (Samanta 1995).
 8 S. Roy et al.
Adaptive response of the TMB to xenobiotics and
insecticides
Variations in the relative toxicity of commonly used
insecticides were observed in the TMB populations from
Jorhat, Assam (Bora and Gurusubramanian 2007; Gurusu-
bramanian and Bora 2008), Darjeeling (Bora et al. 2007),
and the sub-Himalayan Dooars region of India (Roy et al.
2008; Roy et al. 2008c, 2008e, 2009a; Roy, Gurusubrama-
nian, et al. 2009; Roy et al. 2009g; Roy, Gurusubrama-
nian, & Mukhopadhyay 2010b; Roy et al. 2010b, 2011;
Roy and Gurusubramanian 2013). High levels of resis-
tance of the TMB against commonly used insecticides,
particularly endosulfan, quinalphos, cypermethrin and
deltamethrin, have recently been documented (Bora and
Gurusubramanian 2007; Bora et al. 2007; Gurusubrama-
nian and Bora 2008; Roy et al. 2008c, 2008e, 2009a; Roy,
Gurusubramanian, et al. 2009; Roy et al. 2009g; Roy,
Gurusubramanian, & Mukhopadhyay 2010b; Roy et al.
2010b, 2011; Roy and Gurusubramanian 2013; Mukho-
padhyay and Roy 2013). The relative susceptibility values
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(LC50) of the TMB to different insecticides varied by
region. The populations of the Kalchini area of eastern
Dooars in North Bengal showed less susceptibility com-
pared with the fair susceptibility of those from the Dam-
dim and Chulsa areas of Western Dooars to most of the
insecticides commonly in use. The populations of Nagra-
kata and Binnaguri of the central Dooars region repre-
sented an intermediate level of susceptibility (Roy et al.
2008e, 2009g). The LC95 values suggest medium to high
resistance for endosulfan and low to medium resistance for
cypermethrin, lambda cyhalothrin, imidacloprid and quinal-
phos. However, there was not much change for the other
registered insecticides, and they were found to be still
effective at the recommended doses (Roy et al. 2009g).
Selection using sublethal concentrations of endosulfan
and deltamethrin resulted in 4.4- and 5.2-fold increases,
respectively, in the insecticide resistance ratio from the
F1 to F5 generations in the TMB population from a con-
ventional tea plantation (Roy, Gurusubramanian, &
Mukhopadhyay 2010b; Roy and Gurusubramanian 2013).
The biological traits of insecticide-resistant and suscepti-
ble field populations of the TMB revealed that the insecti-
cide-resistant population from the conventional tea
gardens of the Dooars region differed significantly from
the susceptible strain from the organic tea estate of Dar-
jeeling (Roy, Gurusubramanian, & Mukhopadhyay
2010b; Roy et al. 2010b; Roy and Gurusubramanian
2013). In these two populations, the resistant populations
had reduced oviposition period, reduced fecundity, and
prolonged nymphal and total developmental periods. The
resistant forms of the TMB developed certain life cycle
traits to adapt to lower levels of available metabolic
energy and to withstand the stressed conditions resulting
from the repeated exposure to pesticides.
The importance of pigmentation in pesticide resis-
tance in insects is now well established (Srivastava 2004).
Three color variants were observed in males and six in
females of the TMB (Roy et al. 2009f). Three color
variants, both males and females, in the Assam tea planta-
tions have been reported in the same season (Bora and
Gurusubramanian 2007). Blackish forms with a darker
pronotum were frequently encountered in the TMB popu-
lations with high insecticide resistance, and no reddish
orange variant was observed in the Dooars plantations
presumably because of pesticide selection pressure.
Variations in total body lipid content in the TMB pop-
ulations of the Dooars region and the variations in insecti-
cide susceptibility were inversely correlated (Roy et al.
2008c). This correlation emphasized the possible involve-
ment of increased body lipids in furnishing greater toler-
ance to insecticides (Roy et al. 2008c). The increased
activity of defense enzymes such as the GEs, the GSTs,
and the cytochrome P450 mediated mono-oxygenases
(CYPs) was evident in the insecticide exposed TMB pop-
ulations of the Dooars and Terai regions compared with
the unexposed populations in the organic plantations of
Darjeeling (Saha et al. 2012b; Saha and Mukhopadhyay
2013; Saha et al. 2013, Mukhopadhyay et al. 2014). Rela-
tive susceptibility studies conducted by Saha et al.
(2012a) determined that the LC50 values for quinalphos
25 EC were 0.080 ppm for Darjeeling, 43.76 ppm for the
Terai and 214.47 ppm for the Dooars regions. Similarly in
case of cypermethrin 10 EC, the LC50 values were
0.001 ppm for Darjeeling, 3.81 ppm for the Terai and
7.48 ppm for the Dooars regions. The differences in
enzyme activities increased 15.4- to 17.6-fold for the
GEs, 1.8- to 1.9-fold for the GSTs and 2.1- to 2.4-fold for
the CYPs in the Terai and the Dooars populations exposed
to insecticides. A strong correlation was found between
the susceptibility level and the activity of different detoxi-
fying enzymes, particularly with the GEs and cytochrome
P450s (Saha et al. 2012b; Saha and Mukhopadhyay 2013;
Saha et al. 2013).
Exposure to synthetic insecticides such as endosulfan
and deltamethrin drastically changed the egg laying
behavior in the TMB. The majority of the eggs are nor-
mally laid in pluckable shoots and approximately 35% are
inserted into the broken ends of plucked shoots. However,
when the TMB was treated with insecticides such as endo-
sulfan and deltamethrin at sublethal doses, the oviposition
preference of the bug in an insecticide-stressed condition
was reversed. This indication of behavioral resistance
demonstrated by the TMB likely saved more eggs from
insecticide exposure on the tea bush because less of the
spray of these chemicals reaches the nonpluckable portion
of the tea shoots (Roy and Mukhopadhyay 2011).
Therefore, a variety of defense mechanisms, including
enzymatic detoxification systems, physiological tolerance
and behavioral avoidance, protect the TMB from applied
toxic compounds. The feeding activity of the TMB on dif-
ferent host plants may also modulate the level of detoxify-
ing enzyme activity (Saha et al. 2012a). Therefore, this
pest can develop a common strategy to withstand insecti-
cide applications by enhancing the activity of the detoxi-
fying enzymes when exposed to a wide variety of plant
allelochemicals and xenobiotics of diverse groups (Saha
and Mukhopadhyay 2013; Mukhopadhyay and Roy 2013).

Sampling methods
The appropriate methodology for sampling the TMB pop-
ulations is essential, and the sampling method has to be
economically sound and should provide information on
pest abundance with the minimal number of samples.
There are three methods recommended for measuring
TMB populations: (1) counting the average number of
bugs per shoot, (2) counting the percent of affected shoots
that show feeding spots, and (3) collecting shoots from
the baskets of pluckers and counting the number of
infested and uninfested shoots to determine the percent
infestation (Muraleedharan and Selvasundaram 2002). In
Malaysia, a census technique for the TMB along with an
early warning system (EWS) and threshold response sys-
tem was formulated (Wills 1986; Wood and Chung 1989).
Ahmed et al. (1992) developed a computer simulation
model on the population dynamics of H. theivora in
Bangladesh.
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Host plant resistance (HPR)
The importance of HPR in tea has long been presumed
(Watt and Mann 1903) with little understanding of the
defense mechanisms involved (Banerjee 1992). Signifi-
cant morphometric and genetic variability exists among
tea cultivars (Banerjee 1987; Ghosh Hazra 2001), to
which pests react differently (Hazarika et al. 2009). Chi-
nery and hybrid tea jats were found to be more susceptible
to the TMB than “Assam” varieties (Anstead and Ballare
1922; Rau 1940). In Assam, all the tea clones released by
the Tocklai Tea Research Institute were found to be more
or less infested and among them TV1 was the most sus-
ceptible, followed by TV4, TV5, TV6 and TV9 (Das
1984). The clones TV11, TV17, TV21, TV25 and TV26
were reported to be considerably tolerant to this pest
(Somchowdhury et al. 1993). The succulent clones such
as TV1, TV9, TV22, TV25, and TV26 were more suscep-
tible to the TMB (Sundararaju and Sundarababu 1999).
Roy et al. (2009d) screened 28 tea cultivars commonly
cultivated in northeast India and among them TV1, TV12,
TV23, TS653 and TV16 were the most susceptible to
TMB infestation. The clones TV4, TV11, TV28, TV29
and ST449 were less susceptible. The clones TV2, TV9,
TV17, TV18, TV20, TV25, TV26, TV30, Teenali 17,
TS652, TS491, P126, TV7, TV10, TV14, TV19, TV22,
and TS426 were moderately susceptible to TMB infesta-
tion. No clones in northeast India were found to be resis-
tant to the TMB. In south India, the tea clones UPASI 2,
3, 7, 9, 22 and AKK1 were the most susceptible to TMB
infestation. The clones UPASI 10, 12, 14, 15 and 17 were
less susceptible, and the clones UPASI 1, 11, 13 and
ATK1 were moderately susceptible (Sudhakaran 2000).
In Bangladesh, Chowdhury et al. (2008) classified clones
as “fairly resistant,” “resistant,” “susceptible” and “very
susceptible” to the TMB by using a feeding method with
4th instar nymphs for 7 days. Based on these findings, 7
clones, BT1, BT2, BT7, BT8, BT10, BT12 and BT16,
appeared fairly resistant, 8 clones with two biclonal seeds
International Journal of Pest Management 9
and a general seed (BT4, BT5, BT6, BT9, BT13, BT14,
T15 and BT17) showed susceptibility, and two clones,
BT3 and BT11, were very susceptible to the TMB. A lab-
oratory study conducted by Kalita et al. (1997) to investi-
gate the oviposition preference of the TMB on 10
different genotypes of tea revealed that maximum number
of eggs was laid on the clone TV 1, followed by TS 520,
TS 491, TV 28 and TV 9, whereas TV 6 and TV 18 were
the least preferred genotypes under laboratory conditions.
However, the reasons for susceptibility or relative resis-
tance to feeding and oviposition of the TMB are not yet
known. However, TMB infestation leads to a decrease in
phenylalanine ammonia lyase activity and polyphenol
content (Chakraborty and Chakraborty 2001). In addition
to understanding the mechanisms of resistance, the resis-
tance-conferring genes must be identified and incorpo-
rated into the productive cultivars either through
conventional breeding or through genetic engineering
(Mondal 2005). Moreover, describing likely changes in
the proteomes of insects in response to cultivar switching
and insect resistance management will add new dimen-
sions to the HPR programs for the TMB.
Management practices
The perennial and monoculture nature of the tea crop with
the close planting on a large-scale and the applying of suf-
ficient fertilizers and pesticides influences the incidence
of the TMB. The successful management of the TMB is
possible only with early detection and immediate imple-
mentation of management practices. Using multiple con-
trol strategies under IPM, involving cultural, mechanical,
physical, biological and chemical methods, may help
solve the pest problem.
Cultural control
Cultural controls are the first-phase practices and are the
oldest methods that have been used to manage pest popu-
lations. The success of cultural control is dependent on a
detailed knowledge of the bio-ecology of the crop pest,
their natural control and the environmental influences. A
more frequent plucking schedule helped to remove
inserted eggs (Das 1965) and early nymphs on the young
shoots before they could grow large enough to cause more
damage. Hard plucking, black plucking and level off skiff
were effective in cases where there was total or severe
attack. With such styles of harvesting, all the shoots above
the plucking level were removed and the TMB was denied
a food source (Das 1965; Muraleedharan and Selvasun-
daram 2002; Rahman et al. 2006; Roy, Gurusubramanian,
& Mukhopadhyay 2010a). Black plucking and level off
skiff when combined with chemical spraying significantly
decreased the infestation level of the TMB and increase
the crop yield compared with the exclusive use of chemi-
cals (Rahman et al. 2006). In the TMB infested sections,
pruning and skiffing should be conducted from the periph-
ery towards the center, and a few bushes should be
untouched for a day or two in the center to serve as a trap
 10 S. Roy et al.
for adults (Das 1965). These bushes should be pruned/
skiffed after spraying with suitable insecticides. This tech-
nique helps to provide efficient control of the TMB. The
removal of alternate weed hosts of the TMB (Table 1)
from a plantation and the surrounding areas would control
the migration of the pest to other tea field and would keep
the population under control (Das 1965; Roy, Gurusubra-
manian, & Mukhopadhyay, 2010a).
The ecotone between the forest and tea plantation must
be kept clear of weeds and noneconomic plants. The
cleared area needs to be treated with suitable insecticides
to prevent migration of the TMB (Roy, Gurusubramanian,
& Mukhopadhyay 2010a). The TMB prefers moist condi-
tions and mild temperatures. For that reason, this pest is
often observed in large numbers on tea plants under heavy
shade. However, an unshaded condition receiving full sun-
shine is equally detrimental because this enhances attacks
of other pests such as leafhoppers, thrips and red spider
mites. Tea plantations with moderate shade of 60% suffer
less and give optimal crop yields (Rahman et al. 2006).
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Mechanical and physical control methods
There have only been a few attempts to use mechanical
and physical methods in the management of the TMB.
Hand collection of adults and nymphs during minor attacks
is occasionally performed in some gardens, but it should
be started as soon as the signs of damage have been
noticed. The best time for hand collection of the TMB is
the morning and afternoon when the insects are actively
moving on tea bushes. Barrier spraying (a specialized type
of spraying technique where before taking up a spraying
operation of the infested section, about 8 10 rows of tea
bushes at the outer periphery of the section should be
sprayed first and then the operators should start spraying
the remaining part of the section from opposite directions
to prevent migration of the TMB adults to nearby section)
against the TMB was found to be useful (Das 1965). Sus-
ceptible tea clones to the TMB such as TV1 may be used
as a trap crop (Hazarika et al. 2009). Andrews (1914,
1919, 1923) studied the relationship between the TMB and
the soil type and reported that high potash content of the
soil decreased the population of the TMB. He found that
fields having a low ratio of available potash to available
phosphorous were more prone to TMB attack. Ballard
(1921) also emphasized the importance of potassium salts
in the soil as a repellent for the TMB. Ultrasound-based
control may also be a potential component of IPM for the
TMB, which suffered early mortality when exposed to a 20
KHz frequency for 15, 30 and 45 minute per day from the
first instar onward (Borthakur et al. 2011).
Biological control
Conservation and manipulation of natural enemies
Under a classical biocontrol program, attempts would be
made to import exotic natural enemy predators and parasi-
toids for the control of the TMB. However, no attempts
have been recorded. The records of native natural enemies
are scarce, and only scattered information is available on
attempts to apply them through augmentation release pro-
grams. A few natural control agents of the TMB have been
reported (Simmonds 1970; CIBC 1983; Cadou 1994). Watt
and Mann (1903) and Barthakur (2011) reported that a
reduviid bug fed on TMBs. The ant, Crematogaster wroug-
toni Forel (Hymenoptera: Formicidae), was reported to be
a predator of eggs and early instars of the TMB (Ambika
et al. 1979). A survey for egg parasitoids of the TMB from
cocoa plants revealed the prevalence of two parasitoids,
Telenomus sp. and Chaetostricha sp. (Bhat and Srikumar
2013). The reduviids Panthous bimaculatus Dist, Sycanus
collaris Fab and Rihirbus trochantericus luteous Stal were
recorded as effective predators of nymphs and adults of the
TMB on cashew by Bhat et al. (2013). Somchowdhury et
al. (1993) reported that Chrysoperla sp. preyed on the bug,
and under caged conditions, predatory efficiency appeared
to be encouraging. The spider Oxyopes shweta Tikader
was reported as a potential predator of nymphs and adults
(Mukhopadhyay and Sarker 2007; Roy et al. 2006; Kumar
and Mukhopadhyay 2014). The eggs of the TMB were par-
asitized by Erythmelus helopeltidis Gahan (Sudhakaran and
Muraleedharan 1998, 2006) in the southern part of India,
with egg mortality of 52% 83% in laboratory conditions.
Chrysoperla carnea Stephens, Oxyopes sp., Plexippus sp.,
Phidippus sp., Marpissa sp., Mallada sp., Sycanus croceo-
vittatus and praying mantids are other natural enemies of
the TMB in north east India (Das et al. 2010; Barthakur
2011; Borah et al. 2012) (Figure 5). Fungi are the predomi-
nant pathogens found in TMB populations and are unique
in their ability to infect their hosts through the external
cuticle. Gurusubramanian et al. (2009) reported that spray-
ing of the entomopathogen, Beauveria bassiana (Bals.-
Criv.) Vuill, at 3 kg/ ha minimized infestation of the TMB
by 42% 62% compared with the control under field condi-
tions (Figure 6). Field experiments conducted by Ghatak
et al. (2008) in West Bengal to evaluate the efficacy of
biopesticides, viz., B. bassiana, Metarrhizium anisopliae
(Metchnikoff) Sorokin and Verticillium lecanii (Zimmer-
man) Viegas, against the TMB indicated that B. bassiana
had the ability to reduce the TMB infestation by 88%, but
that the other biopesticides were found to be ineffective
against the pest. In a recent study, Bordoloi et al. (2011)
reported that six isolates of the entomopathogenic fungi
Fusarium, Aspergillus flavus Johann Heinrich Friedrich
Link, Cladosporium sp., Curvularia sp., Acremonium and
Trichoderma caused infection in the TMB in the tea
plantations of Assam, India. The mermerthid worm
Hexamermis sp. (Nematoda: Mermithidae) parasitized the
TMB (Mukerji and Roy Choudhuri 1956), and the host
showed no external sign of infestation until the viscera of
the insect was examined.
Botanical pesticides
Botanical pesticides are currently recognized as biodegrad-
able, systemic, eco-friendly and nontoxic to mammals and
are thus considered safe (Isman 2006). Different

Figure 5. (Color online) Natural enemies of Helopeltis Theivora: (A) Oxyopes sp. feeding on adult TMB, (B) Mallada sp. feeding on
nymph of TMB, and (C) Sycanus croceovittatus feeding on adult TMB.
concentrations of aqueous extracts of neem seed kernels
(NKAE) were evaluated against TMB populations with
Downloaded by [Somnath Roy] at 20:10 24 April 2015
antifeedant activity. Additionally, they caused a low hatch-
ing percentage and shortened oviposition and nymphal
periods (Dutta et al. 2013). Neem formulations with differ-
ent azadirachtin contents were evaluated against the TMB
by Roy, Gurusubramanian, and Mukhopadhyay (2010a).
Five different concentrations of azadirachtin were tested,
but only the 50,000-ppm concentration resulted in maxi-
mum (65%) control of the TMB at a higher dilution
(1:200). Because the TMB has sucking type mouthparts,
the chances of ingesting toxicologically active neem (aza-
dirachtin) from the leaf surface are very low. This may be
the principal reason for a low level of control of the TMB
solely with neem formulations. Therefore, the azadirachtin
concentration and its dilutions are the major criteria for
obtaining the desired bioactivity. In addition to neem, a
number of wild plants and weeds available in and around
tea gardens were found to possess insecticidal properties
against the TMB (Table 2). However, most of the informa-
tion on botanicals is restricted to laboratory studies
(Hazarika et al. 2009).
Figure 6. (Color online) Beauveria bassiana infestation on adult TMB.
International Journal of Pest Management 11
Roy et al. (2010a) reported that the water extract of
Clerodendrum viscosum (Verbenaceae), a common weed
in India, showed promise in controlling TMB populations
in the field. Formulations containing azadirachtin combined
with synthetic insecticides, such as endosulfan and delta-
methrin, were reported to be effective for TMB manage-
ment (Roy, Gurusubramanian, & Mukhopadhyay 2010a).
The extract of the same plant species should not be repeat-
edly used for long periods because prolonged exposure
decreases the response of target herbivores (Liu et al.
2005; Hazarika et al. 2009). For organic tea production,
botanicals may play an important role and may be consid-
ered as alternative products for crop protection, particularly
to manage resistance development (Hazarika et al. 2009).
Sex pheromones
Traps using sex pheromones are one of the recent tools
employed in tea pest management. Earlier experiments
conducted by Somchoudhury et al. (1993) and Sudha-
karan (2000) suggested that sex pheromonal activities
exist in the TMB. The attraction of males to female
 Downloaded by [Somnath Roy] at 20:10 24 April 2015

12
Table 2. Plants having anti-insect properties effective against Helopeltis theivora.

Scientific name and family Parts used Extracting medium Mode of anti insect properties Reference

Acorus calamus L. (Acoraceae) Leaves Aqueous Ovicidal, antifeedant Sarmah and Bhola (2011)
Adhatoda vasica Nees Leaves and succulent stems Aqueous, chloroform, petroleum Ovicidal and antifeedant Gurusubramanian et al. (2008a), , Roy
(Acanthaceae) ether, methanol et al. (2009i)
S. Roy et al.

Ageratum conyzoides L. Whole plants Aqueous Ovicidal Roy et al. (2009i)

(Asteraceae)
Allium cepa L. (Amaryllidaceae) Bulb Aqueous Antifeedant, ovicidal Roy et al. (2009i)
Allium sativum L. Bulb Aqueous Antifeedant Roy et al. (2009i)
(Amaryllidaceae)
Amomum subulatum Roxb. Leaves Aqueous Antifeedant Roy et al. (2009i)
(Zingiberaceae)
Annona squamosa L. Leaves and succulent stems Aqueous Antifeedant Gurusubramanian et al. (2008a) and
(Annonaceae) Roy et al. (2009i)
Artemisia vulgaris L. Leaves and succulent stems Aqueous Antifeedant, ovicidal Roy et al. (2009i)
(Asteraceae)
Azadirachta indica A. Juss. Leaves and succulent stems Aqueous Antifeedant activity Sarmah and Bhola (2008)
(Meliaceae)
Chrysanthemum sp. (Asteraceae) Dried flowers Aqueous Antifeedant activity Mamun and Ahmed (2011)
Clerodendron infortunatum L. Leaves and succulent stems Aqueous Antifeedant, ovicidal, Gurusubramanian et al. (2008) and Roy
(Lamiaceae) insecticidal et al. (2008d)
Clerodendron inerme (Linn.) Leaves and succulent stems Aqueous Growth regulating properties, Deka and Saikia (2011) and Deka et al.
Gaertn. (Verbenaceae) antifeedant (2000); Gurusubramanian et al.
(2008a)
Clerodendrum Leaves and succulent stems Aqueous Insecticidal, antifeedant ovicidal Roy et al. (2010a)
viscosum Ventenat
(Verbenaceae)
Datura metel L. (Solanaceae) Leaves and succulent stems Aqueous Insecticidal, antifeedant ovicidal Mamun and Ahmed (2011) Roy et al.
(2008d), and Roy et al. (2009i)
Dentella repens (L.) J.R.Forster Leaves Aqueous Antifeedant Roy et al. (2009i)
(Rubiaceae)
Emblica officinalis Gaertn. Leaves Aqueous Antifeedant Roy et al. (2009i)
(Phyllanthaceae)
Heliotropium indicum L. leaves and flowers Petroleum ether, ethyl acetate Antifeedant activity Dolui and Debnath (2010) and Dolui
(Boraginaceae) and methanol et al. (2012)
Lantana camara L. Leaves and succulent stems Aqueous, chloroform, petroleum Growth regulating properties, Deka and Saikia (2011), Deka et al.
(Verbenaceae) ether and methanol antifeedant, insecticidal (1998), Gurusubramanian et al.
(2008a), and Roy et al. (2009i)
Melia azedarach L. (Meliaceae) Seed, leaves Aqueous Antifeedant, ovicidal Mamun and Ahmed (2011), and
Gurusubramanian et al. (2008)
Momordica charantia L. Leaves and succulent stems Aqueous Antifeedant Roy et al. (2009i)
(Cucurbitaceae)
Nicotiana tabacum L. Leaves and succulent stems Aqueous Antifeedant, ovicidal, Roy et al. (2009i)
(Solanaceae) insecticidal

(continued)

Table 2. (Continued )
Scientific name and family
Pogostemon parviflorus Benth
(Labiatae)
Polygonum hydropiper (L.)
Delabre (Polygonaceae)
Polygonum orientale L.
(Polygonaceae)
Pongamia glabra Vent.
(Fabaceae)
Pongamia pinnata L. (Fabaceae)
Spilanthes calva L. (Asteraceae)
Swietenia mahagoni (L.) Jacq.
(Meliaceae)
Tagetes patula L. (Asteraceae)
Vitex negundo L. (Lamiaceae)
Xanthium strumarium L.
(Asteraceae)
Zingiber officinale Roscoe.
(Zingiberaceae)
Downloaded by [Somnath Roy] at 20:10 24 April 2015
Parts used Extracting medium Mode of anti insect properties Reference
Leaves and succulent stems Chloroform, aqueous extracts Growth inhibitory action, Gogoi et al. (2012), Rahman et al.
Decreased fecundity (2007), Gurusubramanian et al.
(2008a), and Rahman and Nath
(2012)
Leaves and succulent stems Aqueous chloroform petroleum Antifeedant Mamun and Ahmed (2011), Rahman
ether and Nath (2012), and Sarmah and
Bhola (2011)
Leaves and succulent stems Aqueous growth regulating properties, Deka and Singh (2005) and
antifeedant Gurusubramanian et al. (2008a)
Leaves Aqueous Antifeedant Gurusubramanian et al. (2008a)
Seed kernels Aqueous chloroform, petroleum Antifeedant and repellent effects Deka et al. (1998) Deka et al. (2000),
ether and methanol Mamun and Ahmed (2011), and
Gurusubramanian et al. (2008b)
Leaves and flowers Petroleum ether, ethyl acetate Antifeedant activity Dolui and Debnath (2010)
and methanol
Seed kernels Aqueous Insecticidal Mamun and Ahmed (2011)
Whole plants Aqueous Antifeedant, Roy et al. (2009i)
Leaves Aqueous Antifeedant, Roy et al. (2009i)
Leaves and succulent stems Aqueous Ovicidal, antifeedant, Growth Sarmah and Bhola (2008) and Sarmah
regulatory and Bhola (2011)
Rhizome Aqueous antifeedant, ovicidal Roy et al. (2009i)
International Journal of Pest Management
13
 14 S. Roy et al.
pheromones was demonstrated using wind tunnel experi-
ments in the laboratory. Recently, Sachin et al. (2008)
identified the bioactive compound from the thoracic
region of virgin females involved in the pheromonal
attraction of males of the TMB. The behavioral responses
of adult males were mediated by a blend of volatile female
sex pheromone components, (Z)-3 hexenyl acetate and
(E)-2-hexenol, at a ratio of 1:5. This female sex phero-
mone blend may be useful for TMB control and manage-
ment programs.
Use of synthetic chemical insecticides for the control of
the TMB
During the 1940s, application of DDT was practiced to
combat the problem of the TMB. Different pesticides
such as chlordane 10% dust, 50% DDT W.P., Endrex 20
EC, Gammexane 50% W.P., 5% BHC dust, lindane 20%
EC, aldrin, dieldrin, and endrin have been recommended
for controlling tea pests (Glover 1955; Mukerjea 1962). In
the Dooars tea plantation, endosulfan was found to be a
Downloaded by [Somnath Roy] at 20:10 24 April 2015
good standard insecticide, as were DDT and dieldrin
(Mukerjea 1962). After endosulfan was introduced in tea
in northeast India, there was a restriction on the use of
DDT on tea. During the 1970s, eight different types of
chemical insecticides, viz., endosulfan, monocrotophos,
phosalone, Shalimar Tar oil, dimethoate, fenitrothion,
chlorpyriphos and quinalphos, were approved for TMB
management (Banerjee 1973). Synthetic pyrethroids were
first introduced in tea in northeast India in 1982 1983
(Satyanarayana 1982, 1983). Beginning in 2001, several
insecticides, including endosulfan, quinalphos, phospho-
midon, phosalone, acephate, dimethoate, chlorpyriphos,
monocrotophos, oxydemeton methyl, lamda-cyhalothrin,
beta-cyfluthrin, etofenprox, cartap hydrochloride, alpha-
methrin, cypermethrin, deltamethrin, profenfos, thiome-
thoxam, imidacloprid and neem formulations were
recommended for controlling tea pests (Anonymous
2001). However, presently, only a few relatively safer (in
terms of maximum residue limits (MRL) value) insecti-
cides such as deltamethrin, thiomethoxam, bifenthrin, pro-
fenofos, quinalphos and thiacloprid are used for the
management of insect pests of tea.
Rahman et al. (2007) and Roy et al. (2009c; 2010c)
reported on the ovicidal action of different insecticides
against TMB eggs. Profenofos, fenpropathrin, and lamda-
cyhalothrin caused the maximum mortality of eggs. How-
ever, abamectin, azadirachtin, endosulfan, oxydemeton
methyl and acephate had no ovicidal action.
Persistence (PT values) and residual toxicity (LT50
values) of all the commonly used insecticides against the
TMB in northeast India were reported by Roy et al.
(2008). They found that the persistence of neonicotinoids,
synthetic pyrethroids and monocrotophos was between 18
and 28 days, whereas oxydemeton methyl, endosulfan,
and quinalphos persisted for a relatively shorter duration
(7 11 days).
When repeated sprays are necessary, it is important
to rotate insecticides with different modes of action.
Roy et al. 2008 a, b; 2009g; Roy, Gurusubramanian, &
Mukhopadhyay 2010b and Roy et al. 2013 established
that the TMB has developed resistance to endosulfan, qui-
nalphos and deltamethrin insecticides on the Dooars tea
plantation. Under an intensive pesticide application pro-
gram, the level of natural parasitism was very low because
the natural enemies were exposed to excessive amounts of
insecticides (Das et al. 2005). Thus, the failure of insecti-
cides to control the TMB has made management of the
TMB a challenge in most areas of tea production.
Relatively safe insecticides, such as abamectin, cartap
HCl, thiamethoxam and imidacloprid, were suggested for
use by Gurusubramanian et al. (2008a, 2008b, 2009a).
Although a number of chemicals are available for pest
control in tea, for widespread application, the choice of
pesticides is limited to a few for which the MRLs in tea
have been declared by the Food Safety and Standards
Authority of India (FSSAI), the European Union (EU),
the FAO, the Codex Alimentarius Commission and the
US EPA (Environmental Protection Agency).
If solely mediated by synthetic insecticides, it
becomes apparent from the available information that
management of the TMB may become problematic. Con-
trol failures are more common in the tea fields that depend
solely on chemical control strategies (Roy et al. 2009a).
The insecticide-resistant populations of the TMB can be
better managed in the field by integrating various manage-
ment strategies, such as clean cultivation, clipping off of
infested shoots, applying microbial biocontrol agents and
integrating botanicals with synthetic insecticides. How-
ever, the development of location-specific pest manage-
ment modules to combat the TMB through grower
participation would be appropriate in the sustainable man-
agement of this pest.
Future prospects for effective management of the
TMB
The published information on the effects of the TMB on
tea plantations indicates that there are possibilities for
improving the strategies for the effective management of
this sucking pest. Insecticides are often used by growers
on a calendar basis, often leading to resistance, resur-
gence, replacement and residue problems. To improve
this situation, it is important to use all the resources avail-
able in the agroecosystem in a controlled and effective
manner. IPM is an ecosystem-based pest management
strategy that focuses on the longtime control of pests
using a combination of techniques, such as cultural, bio-
logical, and biotechnological methods and habitat manip-
ulation. Chemical pesticides are to be used wisely, only
after monitoring, and under proper guidelines with the
aim to control target pests with no adverse effects on non-
target organisms or the environment. Screening potential
pesticides for disrupting the activity of specific biochemi-
cal sites in the TMB should include targets: transcription
factors belonging to the basic Helix-Loop-Helix (bHLH)
family; antijuvenile hormone (AJH) agents that target JH

biosynthetic enzymes and; G-protein-coupled receptors
(GPCR) for novel methods of pest control.
In the near future, novel biotechnological control strat-
egies could be used that exploit the development of the
thorough knowledge in arthropod genomics and gene
silencing. Integrated biological control methods may be
applied that would involve combined aspects of augmen-
tation and conservation biological control and habitat
manipulation as efficient alternatives to chemical-based
pest management techniques. The success of biological
control strategies depends upon the efficiency of the natu-
ral enemies against the target pest as well as their adapt-
ability, survival and long-term establishment in the
habitat.
The governments or funding agencies of tea-produc-
ing countries should provide adequate financial and logis-
tic support to conduct research on the TMB. This research
should include the novel approaches discussed in this
review and the transfer of technology for better imple-
mentation of the IPM program.
Downloaded by [Somnath Roy] at 20:10 24 April 2015
Acknowledgements
Sincere thanks are due to Mr. Kumar Basnet (Research Fellow,
Department of Zoology, North Bengal University, Siliguri,
India) for providing the photographs of 4th and 5th instar nym-
phals. Thanks are also due to other scientists, and members of
the Entomology Department, Tocklai Tea Research Institute, for
their constant help and support.
Disclosure statement
No potential conflict of interest was reported by the authors.
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