PHYSIOLOGICAL ECOLOGY

Life Table Parameters and Survivorship of Spodoptera exigua

(Lepidoptera: Noctuidae) at Constant Temperatures
AZADEH KARIMI-MALATI,1 YAGHOUB FATHIPOUR,1,2 ALI ASGHAR TALEBI,1
3
AND MOHAMMAD BAZOUBANDI

Environ. Entomol. 43(3): 795803 (2014); DOI: http://dx.doi.org/10.1603/EN11272

ABSTRACT Age-specic life table parameters, survivorship, and extra molting of the beet army-
worm, Spodoptera exigua (Hu bner), were determined at eight constant temperatures of 12, 15, 20, 25,
30, 33, 34, and 36C with a variation of 0.5C on sugar beet leaves. No development was observed at
12 and 36C. The survivorship of overall immature stages was higher at 25C than the other temper-
atures. The highest (0.276 d1) and lowest (0.149 d1) value of the intrinsic rate of increase (r) was
observed at 30 and 20C, respectively. Although the highest value of the net reproductive rate was at
25C (377.7 female offspring at 25C and 127.4 female offspring at 30C), the highest value of r was
at 30C, which indicated the importance of shorter development time (16.9 d at 30C and 27.2 d at
25C) in enhancing r. The relationship between temperature and r was modeled using the Lactin-2
model. The lower temperature threshold, the upper temperature threshold, and optimal temperature
for the r were estimated at 13.1, 32.2, and 34.1C, respectively. The mean generation time decreased
signicantly with increasing temperature, with the longest (35.9 d) and shortest (15.1 d) mean
generation time was calculated at 20 and 33C, respectively.

KEY WORDS beet armyworm, life table parameter, stable stage distribution, extra molting

Among all abiotic factors, temperature has the greatest
effect on arthropods (Ali and Gaylor 1992, Haghani et
al. 2007, Zahiri et al. 2010). Several biological charac-
teristics of insects, such as developmental rate, survi-
vorship, adult life span, sex ratio, fecundity, and fer-
tility, are inuenced by temperature (Roy et al. 2003,
Kheradmand et al. 2007, Kontodimas et al. 2007). In
addition, temperature can affect some physiological
attributes of insects, including extra molting and larval
stadia (Ali and Gaylor 1992).
The beet armyworm, Spodoptera exigua (Hu bner),
is a polyphagous insect that attacks several strategic
crops such as sugar beet, cotton, and soybean. The
damage of the second and third generations is con-
siderable because of the high density of the larvae,
especially in some regions such as California (Ehler
2004). There are multiple factors contributing to the
ineffectual control of S. exigua, even in integrated pest
management programs, including short developmen-
tal time, high fecundity, high vagility, and resistance to
insecticides (Ruberson et al. 1994). Outbreak of S.
exigua is sporadic, but it develops rapidly, and because
older larval instars are difcult to control with insec-
ticides, an early forecasting of the pest is essential for
effective intervention (Saeed et al. 2010). Under-
standing the factors affecting the survival, develop-
1 Department of Entomology, Faculty of Agriculture, Tarbiat Mo-
dares University, P.O. Box 14115-336, Tehran, Iran.
2 Corresponding author, e-mail: fathi@modares.ac.ir.
3 Department of Plant Protection, Khorasan Razavi Agricultural
and Natural Resources Research Center, Mashhad, Iran.
ment, and reproduction of S. exigua may improve the
prediction of its population dynamics and efcacy of
its management programs.
Effects of various host plants on some biological pa-
rameters of S. exigua were previously studied (e.g.,
Meade and Hare 1991; Greenberg et al. 2001; Azidah and
Soan-Azirun 2006; Farahani et al. 2011, 2012; Karimi-
Malati et al. 2012), and few researchers have focused on
development and fecundity of this pest at different con-
stant temperatures on articial diet, cotton, and pigweed
(Butler 1966, Ali and Gaylor 1992). However, published
information about other aspects of the biology of S. ex-
igua such as survival rate and reproductive potential is
inadequate. Furthermore, the life table parameters of S.
exigua at a wide range of temperatures especially on
sugar beet have not been reported yet.
Our study determined the age-specic life table
parameters of S. exigua at eight constant temperatures
on sugar beet. Moreover, the extra molting was studied
at these temperatures. This information might be use-
ful to explain the oscillations in population density as
a function of temperature, and to develop detailed
simulation models to predict phenology of S. exigua
under eld conditions.
Materials and Methods
Rearing Method. The initial population of S. exigua
was originally collected from sugar beet elds of
Khorasan Razavi Agricultural and Natural Resource
Research Center, Mashhad, Iran, during March 2009.

0046-225X/14/07950803$04.00/0 2014 Entomological Society of America

796 ENVIRONMENTAL ENTOMOLOGY
The stock culture of S. exigua was reared on leaves of
sugar beet (Beta vulgaris L.; Renger) and maintained
at 27 1C, 65 5% relative humidity, and a photo-
period of 16:8 (L:D) h in a constant temperature room.
The eggs of the F2 generation were used in this study.
To provide the same aged eggs, adult male and female
moths (10 22 pairs) were kept inside the oviposition
container (12 cm in diameter and 8 cm in height) at
27 1C. The adults were fed on 10% honey solution
impregnated onto cotton wool. After initial oviposi-
tion (few eggs were produced at 27 1C, which were
removed), the containers were transferred to the
growth chambers at eight constant temperatures of 12,
15, 20, 25, 30, 33, 34, and 36C. At rst, development
time was recorded at 12, 15, 20, 25, 30, and 36C. When
the highest development rate was at 30C, and it was
decreased to zero at 36C, another temperature should
be selected between 30 and 36C to obtain the correct
nonlinear point in relationship between temperature
and development rate (the temperature in which the
development rate was lower than that recorded at
30C and higher than zero). Then, the temperature
33C was selected, but the development rate was
higher than the value that was recorded at 30C. Fi-
nally, the temperature 34C was chosen. Before the
experiment, a temperaturerelative humidity data log-
ger (175-H2; Testo, Lenzkirch, Germany) was placed
in the growth chamber to record the temperature
hourly for several days. Therefore, it was possible to
calibrate the growth chamber to have the minimum
variation (1C) during the experiments.
Life Table Parameters. One hundred to three hun-
dred eggs (10 h old) were incubated at the temper-
ature at which they had been laid. The eggs were
checked daily until they hatched or died. The newly
emerged larvae were transferred individually on de-
tached sugar beet leaves in plastic dishes (5 cm di-
ameter and 8 cm height) using a ne camels hair
brush. The lids of dishes had a hole covered with a ne
mesh net for ventilation. The petioles of detached
leaves were inserted into water-soaked cotton, and
fresh leaves were provided daily. The exuviae of head
capsule were used to discriminate the larval molting to
determine extra molting. When larvae developed to
fth instar, some soil (1 cm in depth) was added to the
plastic dishes because the pupation occurs inside the
soil and the pupal cocoon is made of soil. The larval
and pupal survivorship was monitored during the de-
velopmental time and recorded daily. After 3 4 d of
the larvae moving to the soil, the cocoons were cut and
the individuals were sexed at the pupal stage.
Upon emergence, the adult females were paired
with males and transferred to an oviposition container
that was covered on the inside by crumpled and at-
tened paper to provide appropriate oviposition sur-
face. The number of eggs laid was counted daily and
monitoring continued until the death of all adults.
Data Analysis. The life table parameters of S. exigua
at different constant temperatures were determined
by using age (x), survivorship (lx), and female progeny
(mx) according to Careys (1993) equations. Differ-
ences in the net reproductive rate (R0), intrinsic rate
Vol. 43, no. 3
of increase (r), nite rate of increase (), mean gen-
eration time (T), and doubling time (DT) values were
tested for signicance at the examined temperatures
by estimating variances through the jackknife proce-
dure (Efron and Stein 1981). The obtained variables
were tested for normality using the Kolmogorov
Smirnov test before analysis and then subjected to an
analysis of variance (PROC GLM, SAS Institute 2007,
Cary, NC). Moreover, the means were compared by
TukeyKramer procedure (P 0.05).
Models. The linear and nonlinear (Lactin-2) models
were used to t the values of r, which were estimated
at different temperatures for describing the relation-
ship between temperature and r, predicting the values
of r at other temperatures, and determining the lower
temperature threshold (Tmin), upper temperature
threshold (Tmax), and optimal temperature (Topt) for
r (Lactin et al. 1995, Ranjbar-Aghdam et al. 2009). The
r value obtained at 33C was excluded from the linear
model because of it being deviated from linearity. The
Lactin-2 model is as follows:
rT e T e TmTmT/T L
where r(T) is the expected intrinsic rate of increase
estimated by the Lactin-2 model; T is temperature; and
, Tm, T, and L are the model parameters. Estimating
the parameters of the Lactin-2 model and coefcients
of goodness-of-t (SSE, sum of the squared errors;
R2adj, the adjusted coefcient of determination) was
performed by the nonlinear platform of JMP, version
7.0 (SAS Institute 2007).
Results
Survival, Life Expectancy, and Fecundity. The eggs
failed to hatch at 12 and 36C. The age-specic sur-
vivorship (lx) of the beet armyworm at different tem-
peratures are presented in the Fig. 1. The last female
died at the age of 165, 69, 48, 35, 31, and 26 d at tem-
peratures of 15, 20, 25, 30, 33, and 34C, respectively. The
survivorship of overall immature stages was the highest
at 25C compared with the other temperatures tested.
The beet armyworm showed sensitivity to lower and
higher temperatures (15, 33, and 34C) in early ages,
especially at egg hatching, caused a steep decline in lx.
No eggs were produced by the females reared at 15
and 34C. The age-specic fecundity of the beet ar-
myworm throughout the lifespan at 20 33C are dis-
played in the Fig. 2, which revealed that the majority
of the eggs were deposited in the early to middle ages
of females. The age at rst oviposition was 41, 29, 19,
and 16 d at 20, 25, 30, and 33C, respectively. At all
temperatures tested, the mx of the beet armyworm
showed a rapid initial increase to the maximum fol-
lowed by a more clearly dened peak on the second
to fourth day of the oviposition period.
The results revealed that the life expectancy trend of
the beet armyworm was downward at all temperatures
tested. Because of the high mortality of eggs at 15 and
34C, the life expectancy was lower at this stage. After
that, however, the life expectancy showed small changes
in the downward trend during the lifespan (Fig. 3).
June 2014 KARIMI-MALATI ET AL.: LIFE TABLE PARAMETERS OF Spodoptera exigua 797

Fig. 1. Survivorship curves of S. exigua at six constant temperatures.

Life Table Parameters. The life table parameters of
the beet armyworm were signicantly affected by
temperature (Table 1). The R0 varied signicantly
among different temperatures (F 65.87; df 3, 112;
P 0.001). The highest and lowest values of R0 were
observed at 25 and 33C, respectively. The observed r
also differed signicantly (F 150.18; df 3, 117; P
0.0001) according to the pattern of 30 33 25
20C. Moreover, the surprising similarity between the
observed values of r and the estimated ones by the
Lactin-2 nonlinear model at different temperatures is
presented in Fig. 4. The Lactin-2 model tted well
(SSE 0.0032; df 3; R2adj 0.9488) to the obtained
r at different temperatures, indicating that this model
was suitable for describing the relationship between
temperature and r values of S. exigua. The estimated
values of the parameters of this model were 0.0135,
Tm 35.09C, T 0.773, and L 1.192. According
to the estimated values by the Lactin-2 model, the high-
est value of r was 0.312 d1 at 32.2C as Topt. Based on the
output of this model, the minimal and maximal thermal
thresholds that the value of r decreased to zero were 13.1
and 34.1C, respectively. Our laboratory experiments
showed that S. exigua could complete its development at
34C (Karimi-Malati et al. 2014), but the emerged fe-
males were unable to lay fertile eggs.
The showed signicant variation among different
temperatures (F 95.42; df 3, 122; P 0.0001). The
highest observed value of was 1.313 d1 at 30C, and
the lowest value was 1.161 d1 at 20C. In addition, the
signicant difference was observed among the values
of T at different temperatures (F 2016.17; df 3, 111;
P 0.0001). Increasing temperature resulted in a
shorter T (35.9 d at 20C vs. 15.1 d at 33C). Moreover,
798 ENVIRONMENTAL ENTOMOLOGY
Fig. 2. Age-specic fecundity curves of S. exigua at four constant temperatures.
DT was signicantly inuenced by temperature (F larval development time, were combined. The larval
193.62; df 3, 122; P 0.0001), wherein the shortest
DT was 2.5 d at 30C and the longest DT was 4.6 d at
20C.
Development Time and Age- Stage Structure. De-
velopment time and the stable stage distribution for
different life stages of the beet armyworm at 20, 25, 30,
and 33C is summarized in Table 2. The data are not
presented for 15 and 34C because no eggs were laid
at these two temperatures. Variation in egg develop-
ment times was not observed among S. exigua indi-
viduals reared at each temperature because same aged
eggs (eggs laid by the females at the same time) were
used in the experiments. Egg incubation period de-
creased with increasing temperature and varied from
5 d at 20C to 2 d at 30 and 33C. The development time
of the immature stages ranged from 41.63 to 14.5 d at
20 and 33C, respectively.
The results indicate that the stable stage distribu-
tion of S. exigua was independent of temperature (Ta-
ble 2). Proportions of egg, larval, pupal, and adult
stages were approximately constant at 20, 25, 30, and
33C. The durations of different stages are given in
Table 2; therefore, the stable distribution for different
stages could be calculated based on the stable age
distribution. For example, the duration of the egg stage
at 25C was 3 d, so when the stable age distributions
of the rst three days were combined, the egg stage
distribution was 61.08%. Continuously, for larval stage,
the values of the stable age distribution, related to
Vol. 43, no. 3
development time was 14.42 d at 25C; therefore, the
summation of the stable age distributions from day 4 to
day 17 gave the larval stage distribution. It should be
noted that we ignored the variable development time
among individuals and rounded off the development
time to integer and then used the integer age interval to
calculate the stable stage distribution. According to these
calculations, the results showed that the egg stage had
61.0869.51% of the stable age distribution at the tem-
peratures range from 20 to 33C. In fact, the percentage
of all immature stages was 99% at 2033C. At all tem-
peratures tested, the adult stage had the lowest percent-
age of stable age distribution, ranged from 0.08 to 0.52%.
Extra Molting. The current study showed that the
number of larval instars varied at different tempera-
tures. There were ve larval instars at 20, 25, and 30C;
it means that no larvae required more than ve stadia
at the middle range of temperatures (from 20 to 30C)
on sugar beet leaves. At 15 and 34C, all larvae devel-
oped through six instars (100%), while at 33C, 79.2%
of larvae experienced an extra (sixth) stadium. In
addition, some sixth-instar larvae developed to sev-
enth stadium at 33C, but all of them died and could
not complete their larval period.
Discussion
Knowledge of the effect of temperature on demo-
graphic parameters of a given pest is essential for
June 2014 KARIMI-MALATI ET AL.: LIFE TABLE PARAMETERS OF Spodoptera exigua 799

Fig. 3. Life expectancy curves of S. exigua at six constant temperatures.

establishment of an integrated pest management sponse of the beet armyworm to a wide range of
program (Ali and Gaylor 1992, Ranjbar-Aghdam et temperatures to predict its population dynamics in
al. 2009). This study provides the biological re- sugar beet elds.

Table 1. Life table parameters (mean SE) of S. exigua at four constant temperatures

Temperature (C)
Parameters
20 25 30 33
GRR 589.05 30.64a 533.62 22.36a 452.96 19.82b 429.00 23.27b
Fecundity (egg/) 796.42 80.10a 899.10 56.45a 613.16 35.74b 494.32 73.87b
R0 (female offspring) 242.30 19.25b 377.73 20.50a 127.43 7.42c 64.53 7.43c
r (day1) 0.149 0.0028d 0.231 0.0026c 0.277 0.004a 0.263 0.0104b
(day1) 1.161 0.0033c 1.258 0.0034b 1.313 0.0063a 1.304 0.015a
T (day) 35.91 0.22a 25.55 0.15b 17.43 0.18c 15.05 0.19d
DT (day) 4.63 0.089a 3.01 0.036b 2.54 0.045c 2.60 0.012c

Number of eggs laid at 15 and 34C.

Means followed by different letters in each row were signicantly different at P 0.05 (TukeyKramer).
800 ENVIRONMENTAL ENTOMOLOGY Vol. 43, no. 3

Fig. 4. The r values (observed) and r predicted by the Lactin-2 and linear models for S. exigua at four constant
temperatures. The data point 33C has been excluded from the linear regression.

Temperature had a strong effect on lx, R0, and r of
S. exigua. Our ndings revealed that the beet army-
worm was capable of developing across a wide range
of temperatures from 15 to 34C, while the reared
adults did not reproduce at extreme temperatures (15
and 34C). Ali and Gaylor (1992) reported that de-
velopment and lx of the beet armyworm occurs at all
temperatures between 15 and 38C. They emphasized
that no eggs in one mass at 36C and seven masses at
38C were developed. The difference between these
ndings may be related to the difference between the
beet armyworm populations in different geographical
conditions and larval diet.
According to the results of the current study, the
lowest mortality occurred at 25C, which was close to
the ndings reported by Ali and Gaylor (1992) on
articial diet at 26C. The lx of the beet armyworm,
reported in the current study, was somewhat different
from the ndings of Fye and McAda (1972), where the
most efcient rearing was at 30C. They emphasized
that the beet armyworm development was rapid and
the survival rate was 90% on an articial diet at 30C.
The survivorship and life expectancy curves of the
pest at 15, 33, and 34C suggested that the death prob-
ability was high on younger stages compared with old
ones, whereas at moderate temperatures, the mortal-
ity was higher on late ages. The gross reproductive rate
(GRR) was signicantly inuenced by temperature
according to the order of 20 25 30 33C (Table
1). The results of the current study showed that the
highest fecundity occurred at 25C, which was not
signicantly different from that observed at 20C.
However, Fye and McAda (1972) noted that the high-
est fecundity of the beet armyworm was 1,521.9 eggs
at 20C and followed by 25C (874.8 eggs). In addition,
fecundity of the beet armyworm varied from 320.2 to
613.3 eggs on different foliar nitrogen levels at 24.8C
(Al-Zubaidi and Capinera 1984). In the current study,
the maximum R0 occurred at 25C. In fact, the value
of R0 is inuenced by fecundity, survivorship, and sex

Table 2. Duration of development stages (mean SE) and age-stage structure of S. exigua at four constant temperatures

Temperature (C)
Stages
20 25 30 33
Egg
Duration (day) 5.00 0.00a 3.00 0.00b 2.00 0.00c 2.00 0.00c
Distribution (%) 62.68 61.08 63.41 69.51
Larva
Duration (day) 23.20 0.30a 14.42 0.10b 9.82 0.16c 8.43 0.12d
Distribution (%) 36.37 37.58 34.22 27.69
Pupa
Duration (day) 15.30 0.19a 9.78 0.12b 6.14 0.06c 5.22 0.07d
Distribution (%) 0.86 1.17 2.02 2.27
Immature
Duration (day) 41.63 0.36a 27.21 0.16b 16.91 0.20c 14.50 0.16d
Distribution (%) 99.92 99.84 99.64 99.48
Adult
Distribution (%) 0.08 0.16 0.36 0.52

Stable stage distribution could not be estimated at 15 and 34C because no eggs were laid.
Means followed by different letters in each row were signicantly different at P 0.05 (TukeyKramer).
June 2014 KARIMI-MALATI ET AL.: LIFE TABLE PARAMETERS OF Spodoptera exigua
ratio. It seems that the higher survivorship and some-
what higher fecundity is the reason for higher R0 at
25C compared with 20C.
In our study, the recorded range of r for S. exigua
was 0.149 0.276 d1 at 20 33C. According to the
literature, the r value ranged from 0.156 to 0.264 d1
for cabbage and pigweed, respectively, at 26C
(Greenberg et al. 2001). Moreover, Farahani et al.
(2011) determined the range of r of the beet army-
worm to be 0.170 0.264 d1 on different host plants at
26C. However, these differences may be because of
experimental conditions like temperature and host
plant. The parameter r is crucially affected by a wide
range of variables including immature survival, devel-
opmental rate, fecundity, and sex ratio (Jafari et al.
2010). Accordingly, even though the GRR, fecundity,
and R0 values were minimum at 33C, shorter devel-
opmental time led to less x and consequently higher
value of r compared with 20 and 25C. Likewise, re-
search on other species has shown more profound
effect of changes in the developmental rate on r value
than changes in oviposition rate (e.g., Golizadeh et al.
2009a, Jafari et al. 2010, Zahiri et al. 2010, Pakyari et al.
2011, Kouhjani-Gorji et al. 2012). It can be concluded
that the lower survivorship and longer developmental
time led to a lower value of r at 20C, which has been
proved by different scientists (e.g., Haghani et al. 2006,
Zamani et al. 2006, Taghizadeh et al. 2008, Ranjbar-
Aghdam et al. 2009).
In the current study, the results of multiple com-
parisons revealed a signicant difference between r
values at 30 and 33C; however, there was no signi-
cant difference in values at 30 and 33C. Values of
both r and indicate the growth rate and their dif-
ferences among treatments should be consistent. This
coniction may be because of problems caused by
using the jackknife method (Huang and Chi 2013).
The resampling of the jackknife technique results in
unrealistic and meaningless pseudovalues of life table
parameters and overestimation of their variance
(Khanamani et al. 2013). Variance analysis is impor-
tant for determining the variability of experimental
results; hence, the overestimation of variances may
make inconsistent relationships between life table pa-
rameters in different treatments.
Mathematical models are widely used to describe
the reproductive capacity and survival rate of arthro-
pods. The Lactin-2 model was frequently used to pre-
dict the development rate of insects as a function of
temperature (Zahiri et al. 2010). In the current study,
this model was used to simulate reproductive response
of S. exigua to constant temperatures, and it could
predict r successfully because r is strongly correlated
to development rate (Ranjbar-Aghdam et al. 2009,
Golizadeh et al. 2009b, Soufbaf et al. 2010). The ap-
propriate t of the Lactin-2 model to the observed r
allowed us to determine the critical temperatures of
Tmin, Topt, and Tmax as 13.1, 32.2, and 34.1C, respec-
tively. Based on the observed data of this study, the
values of Topt and Tmax estimated by the Lactin-2
model were accurate. However, the Lactin-2 model
underestimated Tmin because our experimental data
801
showed that the females that were reared at 15C did
not lay eggs at this temperature. The inaccuracy might
be due to experimental situations such as unsuccessful
mating at low temperature or other unknown reasons.
It is interesting that females that were mated at 27C
could lay eggs at 15C and even successfully developed
to adulthood at the recent temperature, while the
emerged females were unable to mate successfully or
lay fertile eggs. Another probable reason of the men-
tioned inaccuracy might be because of the intrinsic
weakness of the model in tting data to broad tem-
perature ranges (Roy et al. 2003).
The results of this study indicated that the T at 33C
was approximately half of that estimated at 25C.
Based on Farahani et al. (2011), the value of T varied
from 22.18 to 28.36 d on Brassica napus L. and Gos-
sypium hirsutum L., respectively at 26C. The average
value of T obtained in the current study at 25C (25.55
d) obviously falls within this range.
It seems that the occurrence of ve larval instars is
most common for S. exigua (Al-Zubaidi and Capinera
1984, Meade and Hare 1991, Greenberg et al. 2001,
Farahani et al. 2011). However, other researchers
demonstrated six, seven, or eight stadia for this insect
(Fye and McAda 1972, Ali and Gaylor 1992, Azidah
and Soan-Azirun 2006, Chen et al. 2008). Ali and
Gaylor (1992) found more stadia for the larvae of S.
exigua at extreme temperatures. According to their
results, 3.5% of the larvae reared at 38C (on cotton)
developed to seventh stadium. Furthermore, Azidah
and Soan-Azirun (2006) concluded that the number
of larval instars varies within and between host plants.
The current study showed that 79.2% of the larvae
reared at 33C developed to sixth stadium before pu-
pation. In addition, all larvae at extreme temperatures
(15 and 34C) experienced a supernumerary sixth
stadium. It should be considered that at higher tem-
peratures (33 and 34C), developmental rate was
higher, so the larvae developed to new stadium every
day and the larval development was completed at 7 8
d. In this situation, the size of the fth-instar larvae was
smaller at 33 and 34C compared with 25C (A.K.,
unpublished data). For this reason, extra molting
might be required for the larvae to become larger and
gain critical size. The physiological experiments may
be useful for understanding the hormonal mechanisms
responsible for extra molting at extreme temperatures.
The obtained results conrmed that temperature
strongly affected the biological characteristics of the
beet armyworm. Describing the relationship between
the demographic parameters of the beet armyworm
and temperature would be essential to develop the
population models. This knowledge in conjunction
with complementary semield and led investigations
would contribute to provide a useful pest management
program.
Acknowledgments
This work is a part of the Ph.D. dissertation of the rst
author that was funded by Tarbiat Modares University, Teh-
ran, Iran, which is greatly appreciated. We thank the editor
802 ENVIRONMENTAL ENTOMOLOGY
and anonymous reviewers for their constructive comments
on the manuscript.
References Cited
Ali, A., and M. J. Gaylor. 1992. Effects of temperature and
larval diet on development of the beet armyworm (Lep-
idoptera: Noctuidae). Environ. Entomol. 21: 780 786.
Al-Zubaidi, F. S., and J. L. Capinera. 1984. Utilization of
food and nitrogen by the beet armyworm, Spodoptera
exigua (Hubner) (Lepidoptera: Noctuidae), in relation to
food type and dietary nitrogen levels. Environ. Entomol.
13: 1604 1608.
Azidah, A. A., and M. Sofian-Azirun. 2006. Life history of
Spodoptera exigua (Lepidoptera: Noctuidae) on various
host plants. Bull. Entomol. Res. 96: 613 618.
Butler, G. D. 1966. Development of the beet armyworm and
its parasite Chelonus texanus in relation to temperature. J.
Econ. Entomol. 59: 1324 1327.
Carey, J. R. 1993. Applied demography for biologists with
special emphasis on insects. Oxford University Press, New
York, NY.
Chen, Y., J. R. Ruberson, and D. M. Olson. 2008. Nitrogen
fertilization rate affects feeding, larval performance,
and oviposition preference of the beet armyworm,
Spodoptera exigua, on cotton. Entomol. Exp. Appl. 126:
244 255.
Ehler, L. E. 2004. An evaluation of some natural enemies of
Spodoptera exigua on sugar beet in northern California.
Biocontrol 49: 121135.
Efron, B., and C. Stein. 1981. The jackknife estimate of
variance. Ann. Stat. 9: 586 596.
Farahani, S., B. Naseri, and A. A. Talebi. 2011. Comparative
life table parameters of the beet armyworm, Spodoptera
exigua (Hu bner) (Lep.: Noctuidae) on ve host plants. J.
Entomol. Res. Soc. 13: 91101.
Farahani, S., A. A. Talebi, and Y. Fathipour. 2012. Life table
of Spodoptera exigua (Lepidoptera: Noctuidae) on ve
soybean cultivars. Psyche ID 513824: 17.
Fye, R. E., and W. C. McAda. 1972. Laboratory studies on
the development, longevity, and fecundity of six lepi-
dopterous pests of cotton in Arizona. U.S. Dep. Agric.
Tech. Bull. 1454.
Golizadeh, A., K. Kamali, Y. Fathipour, and H. Abbasipour.
2009a. Effect of temperature on life table parameters
of Plutella xylostella (Lepidoptera: Plutellidae) on two
brassicaceous host plants. J. Asia-Pac. Entomol. 12: 207
212.
Golizadeh, A., K. Kamali, Y. Fathipour, and H. Abbasipour.
2009b. Life table of the diamondback moth, Plutella xy-
lostella (L.) (Lepidoptera: Plutellidae) on ve cultivated
brassicaceous host plants. J. Agric. Sci. Technol. 11: 115
124.
Greenberg, S. M., T. W. Sappington, B. C. Legaspi Jr., T. X.
Liu, and M. Setamou. 2001. Feeding and life history of
Spodoptera exigua (Lepidoptera: Noctuidae) on different
host plants. Ann. Entomol. Soc. Am. 94: 566 575.
Haghani, M., Y. Fathipour, A. A. Talebi, and V. Baniameri.
2006. Comparative demography of Liriomyza sativae
Blanchard (Diptera: Agromyzidae) on cucumber at
seven constant temperatures. Insect Sci. 13: 477 483.
Haghani, M., Y. Fathipour, A. A. Talebi, and V. Baniameri.
2007. Thermal requirement and development of Liri-
omyza sativae (Diptera: Agromyzidae) on cucumber. J.
Econ. Entomol. 100: 350 356.
Huang, Y. B., and H. Chi. 2013. Life tables of Bactrocera
cucurbitae (Diptera: Tephritidae): with an invalidation
Vol. 43, no. 3
of the jackknife technique. J. Appl. Entomol. 137: 327
339.
Jafari, S., Y. Fathipour, F. Faraji, and M. Bagheri. 2010.
Demographic response to constant temperatures in Neo-
seiulus barkeri (Phytoseiidae) fed on Tetranychus urticae
(Tetranychidae). Syst. Appl. Acarol. 15: 8399.
Karimi-Malati, A., Y. Fathipour, A. A. Talebi, and M. Bazou-
bandi. 2012. Comparative life table parameters of beet
armyworm, Spodoptera exigua (Lep.: Noctuidae), on four
commercial sugar beet cultivars. J. Entomol. Soc. Iran 32:
109 124.
Karimi-Malati, A., Y. Fathipour, and A. A. Talebi. 2014. De-
velopment response of Spodoptera exigua to eight con-
stant temperatures: Linear and nonlinear modeling. J.
Asia Pac. Entomol. 17: 349 354. (http://dx.doi.org/
10.1016/j.aspen.2014.03.002).
Khanamani, M., Y. Fathipour, and H. Hajiqanbar. 2013.
Population growth response of Tetranychus urticae to
eggplant quality: application of female age-specic and
age-stage, two-sex life tables. Int. J. Acarol. 39: 638 648.
Kheradmand, K., K. Kamali, Y. Fathipour, and E. Moham-
madi-Goltapeh. 2007. Development, life table and ther-
mal requirement of Tyrophagus putrescentiae (Astigmata:
Acaridae) on mushrooms. J. Stored Prod. Res. 43: 276
281.
Kontodimas, D. C., P. G. Milonas, G. J. Stathas, L. P. Econo-
mou, and N. G. Kavallieratos. 2007. Life table parame-
ters of the pseudococcid predators Nephus includens and
Nephus bisignatus (Coleoptera: Coccinelidae). Eur. J. En-
tomol. 104: 407 415.
Kouhjani-Gorji, M., Y. Fathipour, and K. Kamali. 2012. Life
table parameters of Phytoseius plumifer (Phytoseiidae)
fed on two-spotted spider mite at different constant tem-
peratures. Int. J. Acarol. 38: 377385.
Lactin, D. J., N. J. Holliday, D. L. Johnson, and R. Craigen.
1995. Improved rate model of temperature-dependent
development by arthropods. Environ. Entomol. 24: 68 75.
Meade, T., and J. D. Hare. 1991. Differential performance of
beet armyworm and cabbage looper (Lepidoptera: Noc-
tuidae) larvae on selected Apium graveolens cultivars.
Environ. Entomol. 20: 1636 1644.
Pakyari, H., Y. Fathipour, and A. Enkegaard. 2011. Effect of
temperature on Life table parameters of predatory Thrips
Scolothrips longicornis (Thysanoptera: Thripidae) fed on
twospotted spider mites (Acari: Tetranychidae). J. Econ.
Entomol. 104: 799 805.
Ranjbar-Aghdam, H., Y. Fathipour, D. Kontodimas, G. Rad-
jabi, and M. Rezapanah. 2009. Age-specic life table pa-
rameters and survivorship of an Iranian population of
codling moth (Lep.: Tortricidae) at different constant
temperatures. Ann. Entomol. Soc. Am. 102: 233240.
Roy, M., J. Brodeur, and C. Cloutier. 2003. Effect of tem-
perature on intrinsic rates of natural increase (rm) of
a coccinellid and its spider mite prey. Biocontrol 48:
5772.
Ruberson, J. R., G. A. Herzog, W. R. Lambert, and W. J.
Lewis. 1994. Management of beet armyworm (Lep.:
Noctuidae) in cotton: Role of natural enemies. Fla. En-
tomol. 77: 440 453.
Saeed, S., A. H. Sayyed, and I. Ahmad. 2010. Effect of host
plants on life-history traits of Spodoptera exigua (Lep.:
Noctuidae). J. Pest Sci. 83: 165172.
SAS Institute. 2007. JMP statistics and graphics guide, ver-
sion 7. SAS Institute, Cary, NC.
Soufbaf, M., Y. Fathipour, J. Karimzadeh, and M. P. Zalucki.
2010. Bottom-up effect of different host plants on Plutella
xylostella (Lepidoptera: Plutellidae): a life-table study on
canola. J. Econ. Entomol. 103: 2019 2027.
June 2014 KARIMI-MALATI ET AL.: LIFE TABLE PARAMETERS OF Spodoptera exigua 803

Taghizadeh, R., Y. Fathipour, and K. Kamali. 2008. Inu-
ence of temperature on life-table parameters of Stethorus
gilvifrons (Mulsant) (Coleoptera: Coccinellidae) fed on
Tetranychus urticae Koch. J. Appl. Entomol. 132: 638 645.
Zahiri, B., Y. Fathipour, M. Khanjani, S. Moharramipour, and
M. Zalucki. 2010. Modeling demographic response to
constant temperature in Hypera postica (Coleoptera:
Curculionidae). J. Econ. Entomol. 103: 292301.
Zamani, A. A., A. A. Talebi, Y. Fathipour, and V. Baniameri.
2006. Effect of temperature on biology and population
growth parameters of Aphis gossypii Glover (Hom.: Aphi-
didae) on greenhouse cucumber. J. Appl. Entomol. 130:
453 460.
Received 18 October 2011; accepted 28 March 2014.