Proc. Natl. Acad. Sci., India, Sect. B Biol. Sci.
DOI 10.1007/s40011-016-0750-y
RESEARCH ARTICLE
Host Plant Mediated Susceptibility of Phenacoccus solenopsis
(Tinsley) to Lecanicillium lecanii (Zimmermann) Zare and Gams,
Neem Oil and their Combination
Jaydeep Halder1 Deepak Khushwaha1 A. B. Rai1 K. Nagendran1
B. Singh2
Received: 14 September 2015 / Revised: 25 March 2016 / Accepted: 7 June 2016
 The National Academy of Sciences, India 2016
Abstract Effects of Lecanicillium lecanii alone and in 1:1
combination with neem oil were evaluated against
Phenacoccus solenopsis infesting major vegetables (egg-
plant, tomato, chilli, okra, pointed gourd and cucumber)
and compared with cotton and whitetop weed (Parthenium
hysterophorus Linn.). Marked differences were observed in
the mortality of P. solenopsis amongst the vegetables and
highest percent mortality (66.2 and 90.2 % at 3 and
5 DAT, respectively) was recorded in okra whereas lowest
was in Parthenium (13.9 and 29.9 %) in case of L. lecanii.
Similar trends were also followed in neem oil and their 1:1
combination. Amongst all the vegetables, median lethal
time (LT50) varied from 38.4 to 184.3, 33.7 to 172.8 and
30.6 to 149.2 h in case of L. lecanii, neem oil and their 1:1
combination, respectively. Lowest LT50 value was noted
from of L. lecanii when P. solenopsis fed on okra and their
ascending order of median lethal time was Okra [ Pointed
gourd [ Tomato [ Chilli [ Cotton [ Eggplant [ Cu-
cumber [ Parthenium. Considering the median lethal time
of mealybug reared on okra as base (1), time required for L.
lecanii for 50 % killing of 6 1 day old nymphs of P.
solenopsis was 1.15, 1.38, 2.20, 2.72, 2.75, 2.90 and 4.80
times higher than when fed on pointed gourd, tomato,
chilli, cotton, eggplant, cucumber and Parthenium respec-
tively. From the present study, it is clearly evident that
& Jaydeep Halder
jaydeep.halder@gmail.com
1
Crop Protection Division, ICAR-Indian Institute of
Vegetable Research, P.B. No.01, P.O.- Jakhini
(Shahanshapur), Varanasi, Uttar Pradesh 221305, India
2 or under dosage of insecticide applications is in practice.
ICAR-Indian Institute of Vegetable Research, P.B. No.01,
P.O.- Jakhini (Shahanshapur), Varanasi, Uttar Pradesh
221305, India
strong host mediated variations existed for susceptibility of
P. solenopsis to L. lecanii, neem oil and their combination.
Keywords Median lethal time (LT50)  Neem oil 
Phenacoccus solenopsis  Lecanicillium lecanii
Introduction
Solenopsis mealybug, Phenacoccus solenopsis (Tinsley)
(Pseudococcidae: Homoptera) has a worldwide distribution
with its reported origin in central America [1]. It is an
emerging polyphagous pest and has been observed in
serious proportions on a number of agricultural and horti-
cultural crops and associated weeds [24]. Recently, it has
been observed infesting almost all the vegetable crops
including solanaceous (eggplant, Solanum melongena
Linn; tomato, Solanaum lycopersicon Linn.; chillies,
Capsicum annuum Linn.), cucurbitaceous (cucumber, Cu-
cumis sativus Linn.; pointed gourd, Trichosanthes dioica
Roxb.), malvaceous (okra, Abelmoschus esculentus L.
Moench) vegetables in and around Varanasi region of Uttar
Pradesh, India. Both nymphs and adults cause damage by
sucking the plant sap from the growing points resulting the
infested plants to lose their vitality, remain stunted, leaves
turning yellow, dry up and finally death of plants in case of
heavy infestation. The copious amount of honey dew
excreted by the insects favours the development of the
black sooty mould on leaves and thereby inhibiting the
normal photosynthetic activity of the plants [5].
The indiscriminate and inappropriate use of chemical
insecticides for P. solenopsis management including over
Application of synthetic insecticides frequently leads to
problems like resistance to insecticides, resurgence of
123

target insects, and secondary pest outbreaks in addition to
problems of residues in food and beverages, contamination
of groundwater, adverse effect on human health, and wide
spread killing of non-target organisms [6, 7]. Microbials
and plant-origin insecticides are some of the safe options
due to their target specificity, biodegradability and obvious
safety to the environment. Among the entomopathogenic
fungus, the bioefficacy of white halo fungus, Lecanicillium
(=Verticillium) lecanii against mealybugs are well docu-
mented [8, 9]. Similarly, among the botanicals, neem
(Azadirachta indica A. Juss) is gaining importance due to
its diverse mode of action against wide range of insect
pests across the agricultural and horticultural crops because
of its easy accessibility and lower cost. Earlier study
reported that L. lecanii was most promising and the com-
patibility between the neem oil and L. lecanii against major
sucking pests of vegetables including P. solenopsis
infesting tomato [10]. P. solenopsis, the exotic mealybug
first recorded on cotton from Punjab and Gujarat during
2007 and 2008 [11, 12], respectively has been observed
expanding its host horizon causing severe damage to veg-
etables of diverse families besides weeds including
Parthenium hysterophorus Linn. Looking on its poly-
phagous nature of feeding on the hosts prompted the
authors to find out the host mediated effect on mealybug on
the efficacy of biopesticides alone or in combination with
botanicals, if any.
Material and Methods
Maintenance of P. solenopsis
Initial mealybug P. solenopsis culture was collected from
mealybug infested leaves and twigs of eggplant (Solanum
melongena Linn), okra (Abelmoschus esculentus L.
Moench), tomato (Solanaum lycopersicon Linn.), cucumber
(Cucumis sativus Linn.), chilli (Capsicum annuum Linn.),
pointed gourd (Trichosanthes dioica Roxb.) and whitetop
weed (Parthenium hysterophorus Linn.) from the experi-
mental farm of ICAR-Indian Institute of Vegetable Re-
search, Varanasi (25120 N latitude and 82520 E longitude),
Uttar Pradesh, India. Since the initial occurrence of P.
solenopsis was reported from cotton (Gossypium hirsutum
Linn.), it was also taken as standard to compare the levels of
susceptibility. Each collected sample consisted of varying
LT50 worked out in P:solenopsis feeding on different crops (other than okra) and weed
Relative lethal time
LT50 worked out in P:solenopsis infesting okra
123
J. Halder et al.
number of insects. Mealybugs were released on their
respective hosts under net-house conditions and reared for
three consecutive generations. Only 6 1 day old nymphs
of P. solenopsis were used for the toxicological studies.
Biopesticides
Talc based formulation of entomopathogenic fungi viz.,
Lecanicillium (Verticillium) lecanii (1 9 108 cfu/g) and
neem seed oil (i.e., neem oil) (Himedia Laboratories Pvt.
Ltd., CAS No. 8002-65-1) were taken for the experiment.
Neem oil (1 %) was prepared by dissolving in emulsifying
water containing Triton X-100 (Himedia Laboratories Pvt.
Ltd., CAS No. 9002-93-1). L. lecanii (5 g/l of water) and
neem oil alone and their mixture in 1:1 ratio were sprayed
employing Potters Tower.
Bioassays
Twenty insects of P. solenopsis collected separately from
each host plant were placed in individual petri plates (9 cm
in diameter) and were sprayed with 1 ml of biopesticide
preparations singly and in combinations with neem oil
(1:1) using Potters tower operated at 340 g/cm2 pressure.
Five replications for each treatment per host were main-
tained. The sprayed petri plates containing the treated test
insects were dried for 5 min under fan. Fresh uninfested
and untreated leaves of each host plants were washed in tap
water and shade dried for 30 min and offered to their
respective insects as food. The peduncle of each leaf was
covered with moistened cotton ball to prevent desiccation.
Distilled water spray was served as untreated control. All
the bioassays were done in biocontrol of laboratory under
27 1 C temperature and 70 5 % relative humidity
with 12:12 h light: dark photoperiod. Mortality counts
were taken at 2 and 4 days after treatment (DAT) for the
neem oil and 1:1 mixture of L. lecanii and neem oil where
as in case of L. lecanii data were recorded after 3 and
5 days of treatment. Moribund insects were also considered
as dead. Moreover, regular observations were recorded at
every 12 h interval in each case. Although not truly sus-
ceptible, the okra mealybug population had lowest median
lethal time (LT50 value) values as compared to the rest of
the host plants and hence could serve as baseline (1) in this
study for comparing other mealybug populations. Relative
lethal time was calculated by using the following formula.
1
Host Plant Mediated Susceptibility of Phenacoccus solenopsis (Tinsley) to Lecanicillium

Data Analysis respectively) than their individual effects indicating com-

The mortality data was corrected by Abbotts formula [13]
and analysed by probit analysis [14] with SAS program
(version 9.2). The control mortality in almost all cases, was
below 10 %. The median lethal times (LT50) were deter-
mined and any two values were considered significantly
different if their respective 90 % confidence limits (CL)
did not overlap.
Results and Discussion
Effect of L. lecanii and Its Combination with Neem
Oil (1:1) Against P. solenopsis on Different Host
Plants
Okra
Marked differences were observed between biopesticides
(L. lecanii and neem oil) treated alone and their 1:1 com-
bination for the mortality of P. solenopsis. The white halo
fungus, L. lecanii was found promising against P.
solenopsis as the per cent mortality over control (%MOC)
was 66.2 and 90.2 % at 3 and 5 days after treatment
(DAT), respectively, and took 38.4 h to kill the fifty per-
cent test population (LT50). However, neem oil (1 %) was
found better with % MOC at 2 and 4 DAT and LT50 values
were 61.1 %, 86.5 % and 33.7 h, respectively (Table 1).
When the entomopathogen (L. lecanii) was applied in
combination with neem oil at 1:1 ratio the %MOC were
higher in both the cases (71.3 and 91.9 % at 2 and 4 DAT,
patibility and synergistic activity amongst them. Lowest
LT50 value of 30.68 h was also registered in their 1:1
mixture (Table 2).
Pointed Gourd
Phenacoccus solenopsis when fed on pointed gourd had
highest mortality against L. lecanii (56.1 and 88.3 % at 3
and 5 DAT, respectively) followed by neem oil (53.4 and
85.2 %). Blending of these biopesticides at 1:1 ratio had
shown compatibility and synergistic action as evident from
their highest lethal activity (59.2 and 89.9 % mortality at 2
and 4 DAT, respectively) against P. solenopsis. However,
in comparison with median lethal time (LT50), among the
duo biopesticides, neem oil (1 %) was observed more
effective in controlling the P. solenopsis infesting pointed
gourd and the median lethal time was 39.9 h whereas L.
lecanii had relatively higher (44.2 h) LT50 value. Combi-
nation of these biopesticides at 1:1 ratio registered lowest
LT50 of 33.2 h.
Tomato
Tomato is considered as one of the most preferred host
crop to P. solenopsis [10]. Like okra and pointed gourd, L.
lecanii was able to control this soft-bodied sucking pest in
tomato and gave 59.5 and 86.3 % reduction over control at
3 and 5 DAT and in case of neem oil the corresponding
values were 53.1 and 85.3 at 2 and 4 DAT, respectively
indicating the superiority of neem oil over this EPF.
Median lethal times were also indicating the same trend

Table 1 Effect of L. lecanii and neem oil (1 %) alone and their combination (1:1) against P. solenopsis infesting different vegetables
Treatments Per cent reduction over control
Okra Pointed gourd Tomato Chilli Eggplant Cucumber Cotton P. hysterophorus

L. lecanii
3 DAT 71.3c 59.2a 67.4c 46.2b 25.7a 37.2b 39.5b 17.2b
5 DAT 91.9f 88.3b 86.3d 62.4d 57.2d 52.8e 59.6d 30.6c
Neem oil
2 DAT 61.1a 53.4a 53.1a 40.1a 23.3a 34.6a 32.7a 19.4b
d b d c b d c
4 DAT 86.5 85.2 85.3 59.7 45.8 47.1 55.1 28.1c
L. lecanii ? neem oil (1:1)
2 DAT 66.2b 56.1a 59.5b 44.6b 28.2a 39.5c 37.2b 13.9a
e b de e c c
4 DAT 90.3 89.9 89.1 73.2 52.2 55.2 56.0 29.9c
SEm ()
1.3 1.8 1.6 1.5 2.0 0.9 1.4 1.2
CD (5 %)
2.9 3.9 3.4 3.3 4.3 2.0 3.0 2.6
Means followed by same letter in a column do not differ significantly by DMRT (P=0.05)
DAT days after treatment

123
 J. Halder et al.

Table 2 Median lethal time of L. lecanii and neem oil alone and their combination (1:1) against P. solenopsis infesting different host plants
Biopesticides Heterogenity Regression equation (Y=) LT50 (h) Fiducial limit
2
df v

Okra
L. lecanii 5 0.630 1.537X ? 2.564 38.4 51.8628.46
Neem oil 6 1.467 1.459X ? 2.770 33.7 53.0721.45
L. lecanii ? neem oil (1:1) 6 7.786 2.911X ? 0.677 30.6 39.1723.87
Pointed gourd
L. lecanii 5 0.543 1.852X ? 1.953 44.2 55.2435.29
Neem oil 5 0.973 2.119X ? 1.606 39.9 49.5432.21
L. lecanii ? neem oil (1:1) 5 0.450 1.188X ? 3.194 33.2 55.7319.78
Tomato
L. lecanii 5 0.773 1.897X ? 1.732 52.9 65.4242.78
Neem oil 5 0.984 2.426X ? 0.978 45.5 54.0138.32
L. lecanii ? neem oil (1:1) 5 0.329 2.203X ? 1.455 40.7 49.7133.26
Chilli
L. lecanii 6 1.834 2.342X ? 0.485 84.6 102.2070.05
Neem oil 6 4.136 1.573X ? 2.068 73.2 91.1558.75
L. lecanii ? neem oil (1:1) 6 0.275 1.586X ? 2.185 59.5 75.7046.77
Eggplant
L. lecanii 7 1.263 2.017X ? 0.918 105.7 134.1983.30
Neem oil 8 2.601 3.769X - 2.586 103 115.5491.83
L. lecanii ? neem oil (1:1) 8 5.401 4.184X - 3.256 93.91 103.5185.20
Cucumber
L. lecanii 4 1.256 2.303X ? 0.285 111.5 144.9185.72
Neem oil 4 0.116 1.869X ? 1.153 114.3 165.429.01
L. lecanii ?neem oil (1:1) 5 2.465 2.444X ? 0.378 77.8 92.1165.73
Cotton
L. lecanii 4 2.944 5.898X - 6.909 104.5 116.4793.77
Neem oil 5 1.989 5.163X - 5.397 103.3 115.7792.08
L. lecanii ? neem oil (1:1) 5 8.374 5.074X - 4.703 81.7 89.1874.85
Weed (Parthenium hysterophorus)
L. lecanii 5 8.641 2.518X - 0.704 184.3 273.33124.23
Neem oil 6 7.992 2.636X - 0.897 172.8 232.08128.62
L. lecanii ? neem oil (1:1) 5 0.880 2.464X - 0.357 149.2 230.8996.41

(Table 2). Compatibility and synergistic action were (Table 2). However, blending of this EPF with neem oil
observed at their 1:1 mixture as their mixture had lowest (1:1 ratio) was found compatible and the corresponding
LT50 value of 40.7 h followed by neem oil (45.5) and L. LT50 values were 59.5 h. Similar trend was recorded in per
lecanii (52.9). cent reduction in mortality over control.

Chilli Brinjal

Significant differences were observed in the median lethal
time of L. lecanii and neem oil alone and their 1:1 com-
bination against P. solenopsis feeding on chilli. The
botanical insecticide neem oil was found effective followed
by entomopathogenic fungus L. lecanii when tested against
6 1 day old nymphs of P. solenopsis and their corre-
sponding LT50 values were 73.2 and 84.6 h, respectively
Mealybug reduction over control ranged from 25.7 to
52.2 % at 3 and 5 DAT in case of white halo fungus and
corresponding values for neem oil and 1:1 combination of
L. lecanii and neem oil were 23.445.8 and 28.257.2 %,
respectively (Table 1). The descending order of their tox-
icity is mixture of L. lecanii and neem oil (1:1) [ neem oil
(1 %) [ L. lecanii. In case of median lethal time, lowest

123
Host Plant Mediated Susceptibility of Phenacoccus solenopsis (Tinsley) to Lecanicillium

LT50 value (103 h) was registered against neem oil (1 %)
amongst the duo biopesticides followed by L. lecanii
(105.7 h). Bioactivity of L. lecanii was significantly
enhanced when neem oil was mixed at 1:1 ratio resulting
lowest LT50 value of 93.9 h which was significantly
superior than their individuals.
Cucumber
Phenacoccus solenopsis when fed on cucumber had highest
median lethal time for neem oil (114.3 h) and L. lecanii
(111.5 h) amongst the vegetables. Mixture of neem oil and
L. lecanii had also shown compatible and synergistic
action, and took second highest median lethal time of
77.8 h amongst the vegetables (Table 3).
Cotton
At present the incidence and seriousness of P. solenopsis
on cotton was observed in almost all cotton growing states
of India. So, for a comparative study cotton was also
considered as a host crop. Unlike another malvaceous crop
i.e., okra, the mealybug reduction over control (37.2 and
56.0 % for L. lecanii at 3 and 5 DAT, 32.7 and 55.1 % for
neem oil and 37.2 and 56.0 % for their 1:1 mixture at 2 and
4 DAT, respectively) was significantly lower in cotton. The
LT50 values were 104.5, 103.3 and 81.7 h in case of L.
lecanii, neem oil and 1:1 mixture of L. lecanii and neem
oil, respectively.
Parthenium
During the off-seasons particularly during the months of
MayJune, when none of the above mentioned vegetables are
grown in and around Varanasi, the weed P. hysterophorus
served as an alternate host to this mealybug resulting occur-
rence of this polyphagous mealybug almost throughout the
year in the vegetable ecosystem. Lowest mortality was
recorded amongst all the test cultivated crops as the per cent
reduction in mortality over control in case of L. lecanii was
just 17.2 and 30.6 after 3 and 5 DAT, respectively, and took
highest time (184.3 h) to kill the fifty per cent test population.
Similar trend was also observed in case of neem oil and 1:1
mixture of these biopesticides as their corresponding LT50
values were highest amongst all the test vegetable crops and
cotton. Recently it has been [15] recorded that the exotic
mealybug, P. solenopsis was consistently found attacking
Parthenium plants growing in wastelands of the districts of
Pakistan and anticipated that its year-round occurrence pro-
vides an excellent opportunity for many agricultural pests and
diseases to survive in the unfavorable conditions. Similar
observation was also documented by Arif et al. [16].
Host Mediated Variations
From the Table 3 it is clearly evident that a strong host
mediated variation existed when the biopesticides with
uniform dosages were applied to the same stage of P.
solenopsis collected from different hosts. LT50 values
varied from 38.4 to 184.3, 33.7 to 172.8 and 30.6 to
149.2 h in case of L. lecanii, neem oil and their mixture,
respectively. Amongst all the vegetables under the
study, lowest LT50 value was noted in case of
L. lecanii when P. solenopsis were fed on okra and
their ascending order of median lethal time was as
under Okra [ Pointed gourd [ Tomato [ Chilli [ Cotton
[ Eggplant [ Cucumber [ Parthenium. Since, mealybugs
infesting okra had lowest median lethal time in all the

Table 3 Relative lethal time of L. lecanii alone and its combinations with neem oil (1:1) against P. solenopsis infesting major vegetables and
associated weed
Host plants L. lecanii Neem oil L. lecanii ? neem oil (1:1) CD at 5 % (host wise)

Okra 38.4 (1)a 33.7 (1) 30.6 (1) 1.45

Pointed gourd 44.2 (1.15) 39.9 (1.18) 33.2 (1.09) 3.21
Tomato 52.9 (1.38) 45.5 (1.35) 40.7 (1.33) 3.29
Chilli 84.6 (2.20) 73.2 (2.17) 59.5 (1.95) 5.27
Eggplant 105.7 (2.75) 103.01 (3.05) 93.9 (3.07) 2.24
Cucumber 111.5 (2.90) 114.3 (3.39) 77.8 (2.54) 7.33
Cotton 104.5 (2.72) 103.3 (3.06) 81.7 (2.67) 4.74
Parthenium hysterophorus 184.3 (4.80) 172.8 (5.12) 149.2 (4.88) 5.02
CD at 5 % (treatment wise) 9.73 8.05 8.66
Relative lethal time = LT50 worked out for P. solenopsis feeding on different crops (other than okra) and weed/LT50 worked out in P. solenopsis
infesting okra
a
Figures in the parenthesis are the relative lethal time

123

treatments, comparisons of mealybug on okra was taken as
the base (1). The time required for L. lecanii for 50 %
killing of 6 1 day old nymphs of P. solenopsis was 1.15,
1.38, 2.20, 2.72, 2.75, 2.90 and 4.80 times more when fed
on pointed gourd, tomato, chilli, cotton, eggplant, cucum-
ber and Parthenium respectively, establishing the host
mediated variations. Almost similar trend was observed in
neem oil and 1:1 mixture of L. lecanii and neem oil and the
ascending order of median lethal time for neem oil was
Okra [ Pointed gourd [ Tomato [ Chilli [ Eggplant [
Cotton [ Cucumber [ Parthenium. The present study is in
accordance with the earlier study [17] that the highest
population of third instar mealybug, P. solenopsis
(28.4 mealybugs/6 inch-twig) was recorded on okra fol-
lowed by silvery, tomato, shoeflower and cotton which had
24.6, 23.4, 22.3 and 21.6 mealybugs/6 inch-twig, respec-
tively whereas minimum population of 13.1 mealybugs/6
inch-twig was observed on brinjal. These results can be
compared to the results documented by earlier worker [18],
who reported high density of mealybug on shoe-flower than
the other host plants of P. solenopsis. Host mediated
variation of Spodoptera litura against five insecticides
confirmed that S. litura larvae reared on tobacco were most
tolerant to all the test insecticides whereas those feeding on
sweet potato were most susceptible [19].
Host plant species directly affect the population and
performance of its phytophagous insects and their suscep-
tibility towards management practices. All the dependent
variables, like per cent reduction in mortality of P.
solenopsis over control and median lethal time varied
significantly on different plant species. These variations
might be due to the difference in nutritional quality and
synthesis and nature of allelochemicals present of different
plant species. Amongst all the test cultivable crops, cotton
had relatively higher LT50 values than many of the veg-
etables. Development of resistance in many cotton sucking
pests including P. solenopsis could be a possible reason for
registering the higher median lethal time on the host. In
case of P. hysterophorus, the nutritional quality and pres-
ence of allelochemicals imparting tolerance to P.
solenopsis could be the reason for lower mortality and
higher median lethal time.
Conclusion
Evidence of strong host-mediated interactions were
observed amongst L. lecanii, neem oil and their 1:1 com-
bination and the mealybug, P. solenopsis, infesting major
agricultural and horticultural crops and associated weeds.
This study envisages that there is a need for standardiza-
tions for dosages of these biopesticides for different crops
towards P. solenopsis management.
123
J. Halder et al.
Acknowledgments Authors are thankful to the Director, ICAR-
IIVR, Varanasi, Uttar Pradesh, India for providing the necessary
research facilities for conducting these experiments.
Compliance with Ethical Standards
Conflict of interest All authors declare that they have no conflict of
interest.
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