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Effects of four host plants on biology and food utilization of
the cutworm, Spodoptera litura
Ming Xue1a, Yun-Hong Pang1, Hong-Tao Wang1, Qing-Liang Li1, and Tong-Xian Liu2,3b
1
College of Plant Protection, Shandong Agricultural University, Taian, Shandong 271018, China
2
Department of Entomology, Texas AgriLife Research, Texas A&M University System, Weslaco, Texas 78596-8399,
USA
3
Key Laboratory of Applied Entomology, Northwest A&F University, Yangling, Shaanxi 712100, China
Abstract
Effects of four host plants, tobacco, Chinese cabbage, cowpea and sweet potato, on larval and
pupal development and survival, and longevity and fecundity of adults of Spodoptera litura (F)
(Lepidoptera: Noctuidae), were studied under laboratory conditions (26° C, 60-80% RH), as was
the utilization of the four host plants and adaptation on tobacco. All of the biological parameters
included in the study were affected by the host plants. In a choice test, S. litura females
oviposited most on Chinese cabbage, least on tobacco, and intermediate on cowpea and sweet
potato. S. litura larvae developed differently on the four host plants, from shortest to longest in
the following order: Chinese cabbage, cowpea, sweet potato, and tobacco. Pupal development
was shorter on cowpea than on the other three host plants, and males generally developed longer
than females. More females than males were found among emerged adults, and male adults lived
1-2 d longer than females. Larvae survived best on cowpea (81.6%), followed by Chinese
cabbage (75.5%), then sweet potato (66.1%), and worst on tobacco (49.2%). Pupal survival rates
were relatively high (91.4 - 95.9%) in all four host plant treatments, although that on sweet potato
was lower than those on the other three host plants. Pupal weights on tobacco and sweet potato
were similar, but both were lower than those on Chinese cabbage and cowpea. Generally, male
pupae weighed less than female pupae. Numbers of eggs oviposited by female S. litura were
highest on sweet potato, followed by those on cowpea, Chinese cabbage, and lowest on tobacco.
Relative food consumption rate was highest on sweet potato, followed by that on cowpea,
Chinese cabbage, and lowest on tobacco. In contrast, S. litura larvae that fed on tobacco had
higher efficiency of conversion of digested food, highest efficiency of conversion of ingested
food, and lowest approximate digestibility as compared with larvae that fed on other host plants.
The potential causes for S. litura outbreaks on tobacco are discussed.
Key words: development, food utilization, longevity, oviposition, survival, Chinese cabbage, cowpea, sweet potato,
tobacco
Corresponcence: axueming@sdau.edu.cn, btx-liu@tamu.edu
Received : 8 September 2008 Accepted : 11 April 2009
Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits
unrestricted use, provided that the paper is properly attributed.
ISSN: 1536-2442 | Vol. 10, Number 22
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and many other Asian countries (Shu 1959; and (b) food utilization on the four host plants.
Hill 1975; Shivayogeshwara 1991). S. litura is
also known as the common or tobacco Materials and Methods
cutworm, or the cluster or tobacco caterpillar.
Although it had been a sporadic pest of Host plants
tobacco in northern China for many years, it Four host plants were used in this study,
has been becoming gradually a very important including tobacco (Nicotiana tabacum L.,
insect pest in recent years (Guan and Chen variety ‘NC89’), Chinese cabbage (Brassica
1999; Gao et al. 2004; Qin et al. 2004). It also rapa var. chinensis, variety ‘Shandong Fushen
becomes resistant to many commonly used Baotou’), cowpea (Vigna sinensis L. Walp.
insecticides, particularly pyrethroids and ssp. uniguiculata, variety ‘Zhijiang 28-2’),
carbamates, resulting in failure of effective and sweet potato (Ipomoea batatas L., variety
controls (Wu et al. 1995; Kranthi et al. 2002; ‘Yushu 10’). These plants were selected
Ahmad et al. 2007; Huang and Han 2007). because they are the most important economic
crops in northern China and are primary host
Study of the effects of host plants on the plants of S. litura.
biology of insects is important in
understanding host suitability of plant- The four species of host plants were singly
infesting insect species. There have been a planted in plastic pots (19-cm in diameter and
number of studies on the biological 14 cm in depth) in a greenhouse at Shandong
parameters of S. litura on different host plants Agricultural University at Taian, Shandong,
under different environmental conditions, China, and were maintained insecticide-free.
particularly in India (Patel et al. 1986, 1987), The plants were used when they had 4-5 true
Pakistan (Ahmad et al. 2007), China (Guan leaves.
and Chen 1999; Zhu et al. 2000; Qin et al.
2004; Zhu et al. 2005), Korea (Bae et al. Insects
1997; Bae 1999a,b; Bae and Park 1999), and S. litura larvae were originally collected from
other Asian countries (Etman and Hooper cabbage fields (Brassica oleracea var.
1979; Holloway 1989) where S. litura has capitata L.), and were subsequently reared on
been an important pest on various crops. an artificial diet at the Institute of Pomology,
However, not all of these studied the effects of Guangdong Academy of Agricultural Science,
the same host plants on development, Guangzhou, Guangdong. The artificial diet
survival, pupal weight and oviposition of S. contained the following ingredients: 137 g of
litura under the same environmental corn flour, 10 g of yeast, 37.5 g of soybean
conditions, and none studied the adaptation of flour, 3.5 g of multiple vitamins, 1 g of sorbic
S. litura on tobacco for two generations after acid, 2 g of nipagin, 12.5 g of agar, 0.16 g of
they were reared on an artificial diet. inositol, and 0.15 g cholesterol. Soybean flour
and corn flour were sterilized for 40 min
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conditioned insectary at 26 ± 1⁰ C, 12:12 L:D, readily oviposit on paper stripes (1 cm wide,
and ≈70% RH. S. litura adults were developed 5-10 cm long) in the container. The adults
from the larvae that had been fed with the were monitored daily for mortality and
artificial diet for three generations. Newly oviposition, and number of egg masses
emerged adults were collected and released in oviposited on the paper stripes by each female
each cage at a sex ratio of 1:1. The adults were collected and counted twice daily until
were fed with 10% sugar-water solution the female died.
through a cotton ball in a small plastic
container (3.5 × 1.3 cm). The adults were Food consumption and utilization
allowed to mate in the cage for two days. Newly exuviated sixth instar larvae that had
Eight plants were placed in a screen cage been reared on each of the four host plant
(50×50×50 cm), and two plants from each species for two generations were used in this
species were randomly placed at one of the study. Larvae of approximately the same size
four corners. The plants were adjusted to the were selected and individually placed in small
same height. On the third day, 10 pairs of containers. The larvae were starved for 10 h,
adults (5 females and 5 males) were released and then each larva was individually coded
in each screen cage containing eight plants, and weighed. Twenty larvae were used in
and again, 10% sugar-water solution was each of the four host plant treatments. The
supplied for the adults. The females were larvae were equally divided into a control
allowed to oviposit for 2 days, and the number group and a treatment group. In the control
of egg masses and eggs in each egg mass was group, the 10 larvae and 10 fresh leaves from
recorded for each plant. The experiment was each of the four host plants were individually
replicated six times. weighed, dried in a drier at 80° C, and
weighed again. The dry weights were used as
Larval development and adult the standard for all other treatments. In the
reproduction treatment group, leaves detached from each of
Newly hatched larvae reared on the artificial the four host plants were weighed and
diet were transferred and reared separately in provided to the 10 larvae. The larvae were fed
small containers (3.5×1.3 cm) on each of the with the leaves for 48 h. The larvae were then
four host plants until they reached the fifth starved for 6 h to allow the larvae to defecate.
instar. Newly exuviated fifth instars were The larvae, leaf tissues, and feces in each dish
individually reared in Petri dishes (9.0 × 1.5 were weighed and then dried in a drier at 80°
cm) to avoid cannibalism. Each host plant C. The dried leaf tissues and larvae were
treatment had 100 larvae. The larvae were weighed again. Food utilization rates were
monitored for development and mortality at then calculated based on the formulas of
12-h intervals. In the meantime, the dishes Waldbauer (1968):
were cleaned, and new leaf pieces were
replaced as needed. Before pupation, a few
pieces of paper tissue were placed on the dish
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(17.5 d), cowpea (15.8 d), and shortest on
% Efficiency of conversion of ingested food = Chinese cabbage (13.3 d) (Figure 2A). Of the
[(D-C)/(A-B)]×100 (3) six instars, the first, third and fourth instars
development took significantly longer on
% Efficiency of conversion of digested food = tobacco (F = 58.01 - 91.68; df = 3, 15; p <
[(D-C)/(A-B-E)]×100%) (4) 0.0001) (Figure 2B). In contrast, second instar
development took significantly longer on
Approximate digestibility = [(A-B-E)/(A- sweet potato than on the other three host
B)]×100 (5) plants, and the for the fifth and sixth instars
development time was not significantly
Where A is the weight of dried leaf tissues in different for tobacco and sweet potato (Figure
the control, B is the weight of the dried leaf 2B). Pupal development times on Chinese
tissue in each treatment, C is the weight of cabbage (10.9 d), cowpea (10.1 d) and sweet
dried larvae in the control, D is the weight of potato (10.1 d) were not significantly
dried larvae in each treatment, and E is the different, and were longer than on cowpea
weight of dried feces in each treatment. (9.5 d) (F = 6.84; df = 3, 15; p < 0.05) (Figure
2A).
Data analysis
The life history parameters of S. litura were Larval and pupal survival
analyzed using one-way and factorial The survival rates of S. litura larvae varied on
ANOVA (SAS Institute 2008). Means the four host plants (Figure 3). The overall,
associated with host plants for each variable accumulated survival rates of all larval stages
were separated using the least significant on the four host plants differed significantly
difference test when significant values were (F = 26.43; df = 3, 15; p < 0.05) (Figure 3A)
obtained. and was lowest on tobacco (49.0%), followed
by that on sweet potato (66.2%), on Chinese
Results cabbage (75.4%), and highest on cowpea
(81.7%). Of the six larval stages, the survival
Host plant preference for oviposition rates of S. litura were significantly different in
In the choice test, numbers of egg masses the first four instars (F = 4.03 - 42.27; df = 3,
oviposited by S. litura females on the four 15; p = 0.0339 - 0.0001), but not in the two
host plants differed significantly (F = 17.73; oldest ones (F = 0.83 and 2.62; df = 3, 15; p =
df = 3, 23; p = 0.002) (Figure 1). S. litura 0.0987 and 0.5018) (Figure 3B). Different
oviposited the most on Chinese cabbage (13.3 larval instars responded differently on each of
egg masses, 36.7%), followed by sweet potato the four host plants. The survival rates of the
first instar were highest on sweet potato
(9.3 egg masses, 26.7%) and cowpea (9.0 egg
masses, 24.8%), and the least on tobacco (4.7 (99.0%) and lowest on tobacco (80.1%), with
egg masses, 12.8%). intermediates on Chinese cabbage (90.0%)
and cowpea (91.7%). The survival rates of the
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Figure 1. Egg masses per female Spodoptera litura on four host plants. Host plants: CC - Chinese cabbage, CP -
cowpea, SP - sweet potato, and TO - tobacco. The same letters over the four bars in each figure indicate that the
means are not significantly different at p < 0.05 (Least Significance Difference test, SAS Institute 2008). High quality
figures are available online.
Figure 2. Development of Spodoptera litura larvae on four host plants. Host plants: CC - Chinese cabbage, CP -
cowpea, SP - sweet potato, and TO - tobacco. The same letters over the paired-bars in each figure indicate that the
means are not significantly different at p < 0.05 (Least Significance Difference test, SAS Institute 2008). High quality
figures are available online.
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Figure 3. Accumulated survivals (A) and stage-specific survivals (B) of Spodoptera litura on four host plants. Host
plants: CC - Chinese cabbage, CP - cowpea, SP - sweet potato, and TO - tobacco. The same letters over the four
bars in each figure indicate that the means are not significantly different at p < 0.05 (Least Significance Difference test,
SAS Institute 2008). High quality figures are available online.
Table 1. Effects of four host plants on pupae and adults of Spodoptera litura.
Pupal weight Sex ratio Longevity days ± SE
g/pupa ± SE (female:
male)
Host plants Female Male Female Male
Chinese cabbage 0.362±0.003a 0.354±0.002a 1:0.64b 6.6±0.2b 7.4±0.2b
Cowpea 0.346±0.008a 0.310±0.007b 1:0.71a 7.7±0.6a 8.8±0.7a
Sweet potato 0.341±0.013ab 0.318±0.008b 1:0.75a 6.8±0.8b 8.8±0.6a
Tobacco 0.320±0.013b 0.317±0.011b 1:0.64 b 7.2±0.4a 8.2±0.4ab
F3,15 19.76 38.69 19.52 6.15 10.06
P 0.0001 0.0001 0.0001 0.089 0.0014
The means (±SE) in the same column followed by the same letters are not significantly different at P < 0.001 (LSD, SAS
Institute 2008).
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rates of the third instars were similar on
Chinese cabbage (100%) and cowpea Sex ratio, adult longevity and oviposition
(95.5%), which were higher than those on Sex ratios were biased, and more female
sweet potato (86.8%) and tobacco (83.8%). adults emerged than male adults when their
The fourth instars survived less on tobacco larvae were fed with the four host plants (F =
(90.3%) than on the other three host plants 19.52; df = 3, 15; p = 0.0001) (Table 1). Of
(98.6 - 100%). The survival rates for prepupae the six treatments, male ratios were higher on
were not significantly different (F = 0.93; df = cowpea and sweet potato than on the other
3, 15; p > 0.05), but those for pupae were (F = two host plants. The longevities of both
4.56; df = 3, 15; p < 0.05). female and male S. litura adults were also
significantly affected by the host plants on
Pupal weight which their larvae fed (female: F = 6.15; df =
Pupal weights differed significantly depend- 3, 15; p = 0.0089; male: F = 10.6; df = 3, 15;
ing on the host plants on which the larvae p = 0.0014). Numbers of egg masses and total
were fed and differed significantly between eggs oviposited by S. litura females on the
females and males when they fed on the same four host plants differed significantly (egg
host plants and when larvae fed on different masses: F = 16.61; df = 3, 15; p = 0.0001;
host plants (p < 0.005) (Table 1). The female total eggs: F = 31.13; df = 3, 15; p = 0.0001)
pupae on Chinese cabbage were heaviest, (Figure 4). S. litura oviposited similar
followed by those on cowpea and sweet numbers of egg masses and eggs per female
potato, and lightest on tobacco, and the on cowpea, sweet potato, and Chinese
cabbage, but less on tobacco than on the other
Figure 4. Egg masses and total eggs per female Spodoptera litura on four host plants. Host plants: CC - Chinese cabbage,
CP - cowpea, SP - sweet potato, and TO - tobacco. The same letters over the four bars in each figure indicate that the
means are not significantly different at p < 0.05 (Least Significance Difference test, SAS Institute 2008). High quality figures
are available online.
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(233.6 eggs/egg mass). 3, 39; p < 0.001) and were higher on Chinese
cabbage and cowpea than on sweet potato and
Food consumption and utilization tobacco.
Food consumption and conversions of in-
gested and digested food by S. litura larvae Discussion
varied considerably among the four host
plants that the larvae consumed (Table 2). The The data clearly show that S. litura performed
relative growth rates on tobacco (0.43), differently in oviposition, larval and pupal
cowpea (0.43) and Chinese cabbage (0.40) development and survival, pupal weight, and
were higher than that on sweet potato (0.32) oviposition of emerged females when Chinese
(F = 8.050; df = 3, 39; p < 0.001). The relative cabbage, cowpea, sweet potato and tobacco
consumption rates were highest when the were offered as the food plants for their
larvae fed on sweet potato (3.90), followed by larvae. These results are supported by a
that on cowpea (3.16), then on Chinese number of studies, although direct comparison
cabbage (2.28), and the lowest on tobacco of these data can be difficult because different
(1.51) (F = 56.19; df = 3, 39; p < 0.001). The host plants and environmental conditions were
efficiency of conversion of ingested food was used in these studies. Although the same
highest on tobacco (29.75), followed by that insect was used, it differed in origins, and it
on Chinese cabbage (17.85), then that on could be different strains or biotypes.
cowpea (14.04), and lowest on sweet potato However, in a few studies, the same host
(8.34) (F = 74.59; df = 3, 39; p < 0.001). The plants (one or two) were used, including
efficiency of conversion of digested food was cowpea (Qin et al. 2004; Zhu et al. 2005) and
higher when the larvae fed on tobacco than Chinese cabbage (Zhang et al. 1997). Qin et
when fed on the other three host plants (F = al. (2004) used the same variety of cowpea but
18.73; df = 3, 39; p < 0.001).. However, the found that the developmental time of S. litura
Table 2. Nutritional indices of Spodoptera litura larvae feeding on four host plants.
Rate ± SE
Chinese Cowpea Sweet Tobacco F3,39 P
Category cabbage potato
Relative growth rate 0.40±0.02a 0.43±0.02a 0.32±0.02b 0.43±0.01a 8.05 0.0003
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This difference (5.7 d) could be caused by These results show that larval food directly
higher temperature (28.1° C) or different affects pupal size and weight, and the female
cowpea variety from what was used here (26° pupae were heavier than male pupae on all
C; ‘Zhijinag 28-2’). four host plants (Table 1). The pupal weights
of S. litura in this study were 0.32 to 0.36 g,
Larval development of S. litura varied greatly which were generally within a wide range on
depending on host plants and temperature, and various host plants, from 0.28 g on sweet
the development was prolonged under low or potato to 0.40 g on perilla and cowpea (Bae
high temperatures (Zhu et al. 2000; Chen et al. and Park 1999; Qin et al. 2004). Pupal
2002; Seema et al. 2004). For example, on weights were not found to be significantly
tobacco, larval development can range from different among different temperature regimes
19.3 d at 26° C (Chen et al. 2002), 23.2 d in (Seema et al. 2004), but 27° C was considered
this study at 26° C, to as long as 30 d at 23° C the optimum temperature for larval and pupal
(Rattan and Nayak 1963). Similarly, on growth and development. Bae and Park (1999)
cowpea, larval development ranged from 10.1 found that pupal weight tended to be 3 - 13%
d at 28° C (Zhu et al. 2005) to 15.8 d at 26° C lower with increasing temperature from 24° C,
in the present study. to 28° C, then to 32° C.
It has been reported that pupal development In the present data, more than 91% of S. litura
was not affected by host plants on which their pupae successfully developed to adults. In
larvae fed (Patel et al. 1986). However, our contrast, Bae and Park (1999) reported
results show that pupae developed faster on significantly low emergence rates on four
cowpea than on Chinese cabbage, sweet foods. They found that only 31.4% of pupae
potato and tobacco, although the difference developed to adults at 24° C and 88.6% at 30°
was <1 d longer on sweet potato, 1 d longer C, and their emergence rate increased 1.1 to
on tobacco, and 1.5 d longer on Chinese 2.8 fold with temperature increase from 24° C
cabbage. Bae and Park et al. (1999) found that to 32° C.
temperature plays a vital role on pupal
development. Bae and Park (1999) reported Numbers of eggs of S. litura oviposited by the
that the mean pupal developmental duration females from the larvae that fed on the four
was as long as 13.8 d at 24° C on perilla to as host plants were generally within the range as
short as 7.4 d at 32° C on soybean. reported on various host plants (Figure 3). In
the literature, oviposition by females varied
Larval survival or pupation rate of S. litura greatly on different hosts under different
varied greatly on different host plants, ranging environmental conditions (Patel et al. 1986;
from 100% on Ricinus communis (Patel et al. Bae and Park 1999). Numbers of eggs laid by
1987) to 49.0% on tobacco in this study. Bae a single female ranged from 935 on soybean
and Park (1999) found that pupation rates (Bae and Park 1999) to 3,467 on cotton (Patel
were positively correlated with high et al. 1986). In contrast, Bae and Park (1999)
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its "ideal" growth rate even with foods of
In the present study, male adults generally different quality. Zhu et al. (2005) found that
lived longer (7.4 - 8.8 d) than females (6.6 - S. litura larvae did not prefer feeding on
7.7 d), differing on different host plants (Table banana leaves and had lower relative growth
1). Similar results were reported by Bae and rate, relative consumption rate, and
Park (1999), although the differences were approximate digestibility, but it had a
generally less than 1 d (0.25 - 1.0 d). significantly higher efficiency of conversion
However, Patel et al. (1986) found that on of ingested food and an extremely higher rate
cotton, male adults lived 6.3 d as compared of efficiency of conversion of digested food,
with 12.3 d for female adults. It has been indicating that the larvae are capable of
found that adult longevity became shorter as compensating by more efficiently utilizing
the temperature increased (Bae and Park their limited banana leaf tissues than other
1999). In the present study, the sex ratio was host plants. The digestion rate is affected by
biased, and more females emerged than males the enzyme activities of various host plants,
with sex ratios of 1.0:0.64 - 0.75. This was including trehalase, invertase, and others. In
supported by Seema et al. (2004). Differences practice, however, it can be quite difficult to
in pupal survival, adult sex ratio, longevity, ascertain "cause" and "effect" responses with
and fecundity may also be affected by efficiency parameters. Does the insect eat
temperature and other environmental more because digestibility is low, or is
conditions (Zhu et al. 2000; Chen et al. 2002; digestibility low because the insect is eating
Seema et al. 2004). more? Efficiency parameters are so closely
related physiologically that determination of
Our data show that all nutritional indices "cause" and "effect" is not a trivial matter.
varied when S. litura fed on the four host Factors contributing to such changes are still
plants. Food conversion efficiencies on largely unknown, but may include shifts in
different host plants vary considerably by S. food selection, digestive physiology,
litura larvae (Balasubramanian et al. 1985) metabolic rates, and body composition.
and by insects in general (Scriber and Slansky Understanding of these basic principles of
1991; Slansky and Scriber 1985). The current nutritional ecology can enhance our
data show that S. litura had similar relative appreciation of insects’ adaptation to new
growth rates on Chinese cabbage, cowpea, food resources.
and tobacco, but had the lowest relative
consumption rate when feeding on tobacco As this study found that S. litura dose not
compared with those for the other three host prefer feeding on tobacco, why has it been
plants. However, the larvae were more able to cause severe damage on tobacco in
efficiently converting tobacco tissues into China? Our data indicate that S. litura was
their biomass than other plant tissues as able to adapt on tobacco in a relatively short
shown by larvae fed on tobacco having the period of time (Xue, unpublished data). For
lowest approximate digestiblity and the example, comparison of the development,
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explain the facts that S. litura could adapt on four host plants were ranked as Chinese
tobacco and quickly become a severe pest. cabbage > cowpea > sweet potato > tobacco.
The outbreaks of S. litura can be affected by Hence, the present study has shown the
many biotic and abiotic factors. Gao et al. suitability of selected host plants for the
(2004) identified several factors that may development, longevity, and survival of S.
cause outbreaks of S. litura in northern China. litura. These findings will help to understand
Those factors include increasing acreage of the biology of this particular pest and could
tobacco and many preferred crops (mainly help in its management and control,
vegetables), which provides abundant food particularly on tobacco. Therefore, future
sources; expanding of protected cultivations, studies should focus on testing a wider range
which provides suitable sites for of host plant species for the development of S.
overwintering; more mild winter and warmer litura, and assessment of the chemical
spring, which enable the pest to occur earlier components of the host plant species would
and build up higher spring populations in the help to better understand the mechanism of
first generation; high temperature and less host suitability.
rainfall in summer; and misuse of pesticides
that cause resistance to insecticides and less Acknowledgements
natural enemies. Monoculture could be We would like to thank Y. Zhang, L. Q. Wei
another factor because it is more favorable for and L. L. Li for technical assistance, and the
the pest, and irrigation may also contribute to Institute of Pomology, Guangdong Academy
the outbreaks of S. litura in China. The of Agriculture Science (Guangzhou,
occurrence of S. litura is generally Guangdong, China) for providing the insects
synchronous with the rapid growing period of for this study. This work was financially
tobacco, especially during summer, providing supported by The State Tobacco Monopoly
plenty of sources for oviposition and larval Administration, China Tobacco Corporation
feeding. In India, Pandey (1977) considered a (2003579), and The Nature Science
good rainfall after a long period of drought as Foundation of China (30671379).
the key factor causing S. litura outbreaks. S.
litura have also become resistant to most References
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