Open Agriculture 2024; 9: 20220254

Research Article

Danar Dono*, Retno Wulansari, Yusup Hidayat, Neneng Sri Widayani

Biology of Spodoptera frugiperda (Lepidoptera:

Noctuidae) on different types of plants feeds:
Potency as a pest on various agricultural plants
https://doi.org/10.1515/opag-2022-0254 the nutritional indices of several feed plant species tested
received July 8, 2023; accepted January 3, 2024 provided information that broccoli could be the most sui-
Abstract: Spodoptera frugiperda is a new invasive and table host compared to other plants tested when there was
highly polyphagous pest that attacks corn in Indonesia. no corn in the field.
The availability of abundant plant species allows pests to Keywords: fall armyworm, insect biology, life table, nutri-
switch to other host plants to maintain their population. tional indices, host suitability
The aim of this research is to examine the development,
reproduction, nutritional indices, and life table of S. frugi-
perda in several plant species. The plants tested were corn,
rice, broccoli, oil palm, and baby corn as controls. Ten 1 Introduction
individual insects were used and the experiments were
repeated five times for each plant species. The test results Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae)
show that different types of plant feed affect the develop- was the first reported pest in 1797 in Georgia, America [1].
ment time, imago life span, fecundity, and fertility of S. Furthermore, this pest in 2016 was first reported in West
frugiperda. Baby corn fruit and broccoli had higher net and Central Africa [2]. Also, S. frugiperda populations were
reproduction value (R0), intrinsic growth rate (r), gross discovered in 2018 in many locations in India [3] and South-
reproduction rate (GRR), shorter mean generation period east Asia in 2019 [4]. In early 2019, it was reported to be
(T), and population doubling time (DT) than corn and rice discovered in Indonesia [5].
leaves. On oil palm leaf feed, no population parameters The migration ability of S. frugiperda imago is high up
could be determined because no larvae developed into to 1,600 km in 30 h [6]. Hence, it spread rapidly to several
adults and had the lowest nutritional indices parameters, areas in Indonesia. For example, it was reported in West
so that, oil palm could not be exploited as a host plant. Also, Sumatra in March 2019 [5] and in May 2019 S. frugiperda
was confirmed in Lampung [7], and in in June 2019,
Maharani et al. [8] reported that this insect was spread
 at Bandung, Garut, and Sumedang district. The pests attack
* Corresponding author: Danar Dono, Department of Plant Pests and
corn plants from the vegetative to generative phases.
Diseases, Faculty of Agriculture, Universitas Padjadjaran, Bandung,
45363, West Java, Indonesia; Pesticide and Environmental Toxicology
S. frugiperda is highly polyphagous, causing economic
Laboratory, Division of Centre for Product Development and Partnership impacts on various plants [9]. Moreover, 353 different host
Study (Puspromit), Faculty of Agriculture, Universitas Padjadjaran, plant species have been reported from 76 plant families
Bandung, 45363, West Java, Indonesia, e-mail: [email protected] based on a thorough literature review, especially the Poa-
Retno Wulansari: Department of Plant Pests and Diseases, Faculty of ceae, Asteraceae, and Fabaceae families [10], as well as the
Agriculture, Universitas Padjadjaran, Bandung, 45363, West Java,
Brassicaceae family as other host [11]. The availability of
Indonesia
Yusup Hidayat: Department of Plant Pests and Diseases, Faculty of abundant hosts makes S. frugiperda populations able to
Agriculture, Universitas Padjadjaran, Bandung, 45363, West Java, create new food preferences on various plants.
Indonesia; Pesticide and Environmental Toxicology Laboratory, Division In line with Subramanian and Mohankumar [12], the
of Centre for Product Development and Partnership Study (Puspromit), ability of insects to survive on diverse host plants is an
Faculty of Agriculture, Universitas Padjadjaran, Bandung, 45363, West
adaptive mechanism for survival in ecosystems. However,
Java, Indonesia
Neneng Sri Widayani: Agricultural Science Program, Faculty of
the availability and quality of host plants play an impor-
Agriculture, Universitas Padjadjaran, Bandung, 45363, West Java, tant role in pest population dynamics by influencing larval
Indonesia and imago stage performance [13]. Furthermore, according

Open Access. © 2024 the author(s), published by De Gruyter. This work is licensed under the Creative Commons Attribution 4.0 International License.
2  Danar Dono et al.

to Chapman [14], the amount and quality of food consumed conditions. Moreover, the plants were watered regularly, and
by insects affects development, reproduction, and life span. then fertilization was carried out 7 days after planting with a
Based on the pest management perspective, the life table dose of NPK (15:15:15) fertilizer of 3 g per plant for corn and
is essential to determine the most vulnerable pest stage for broccoli plants, as well as fertilization was carried out 14 days
pest control [15]. Therefore, this research was conducted to after transplanting for rice plants.
examine the development, reproduction, nutritional indices, Replanting was carried out once a week to obtain uni-
and life table of S. frugiperda in several plant species. In form and sufficient plants to feed the larvae. Plants can be
addition, S. frugiperda has several alternative hosts that can used as feed after they have more than 5 leaves or are
be utilized apart from its primary host. As an implication, we more than 2 months old. Meanwhile, oil palm leaves
can ascertain the potential of S. frugiperda as a pest on var- were obtained from young plants in Jatinangor, West Java
ious plants (food sources) used in this study.

2.3 Effect of feed types on biology of

2 Materials and methods S. frugiperda

2.1 Rearing of test insect Larvae that emerge from newly hatched eggs (<24 h) were
selected to be placed in plastic cups as many as 50 larvae
S. frugiperda larvae were obtained from corn plantations for each feed type. The larvae were placed separately in
in Jatinangor, Sumedang, West Java. Indonesia. The larvae plastic cups (diameter of 2.5 cm, height of 4 cm) with leaves
were reared at the Pesticide and Environmental Toxicology of corn, rice, broccoli, oil palm, and baby corn fruit as
Laboratory, Department of Plant Pests and Diseases, Faculty control. Feed leaves and baby corn fruits were replaced
of Agriculture, Universitas Padjadjaran, West Java, Indonesia. daily with fresh ones.
Furthermore, the larvae were kept in plastic boxes measuring Observations were made every day to determine the
34 × 28 × 7 cm. Early instar larvae kept in plastic boxes were mortality and development time of larvae. In addition, the
fed with baby corn fruit (Zea mays). Before pupation, the larvae length of larval development was observed by recording
were transferred to a plastic container with a line of paper and the time required for S. frugiperda larvae to develop from a
given sawdust as the medium for pupation. Furthermore, the certain instar to the next, marked by the molting of the
pupae that had been formed were transferred to the cage larval cuticle. After the larvae became pupae, observations
(measuring 44.5 × 44.5 × 49.5 cm) until they became an imago. were made, including the development time and weight of
The imagos were fed 10% liquid honey absorbed on a pupae, normal and abnormal conditions, as well as mor-
lump of cotton. Afterwards, pesticide-free corn leaves were tality. Weight of pupae was measured on the third day
put in bottles filled with water and placed in plastic cages after pupation using analytical balance.
as a place to lay eggs. The eggs laid by imago on corn leaves The imago that emerged from pupae were paired in a
were collected daily and placed in a ventilated plastic box cage (diameter of 13.5 cm and height of 13 cm) where a 10%
measuring 10 × 9 × 4.5 cm, lined with paper at the bottom. honey solution was absorbed into the cotton as food of imago.
Moreover, insects were maintained every day, hence, the Furthermore, the corn leaves, rice leaves, and broccoli leaves
larvae were available for testing. are placed in the cage (according to the larval feed) to laid
eggs. The number of eggs laid by each female and mortality of
imagos were recorded daily. Moreover, dead female imagos
2.2 Feed plant cultivation were dissected to reveal ovaries. A lateral incision was made
in the abdomen following the midline of the thorax from the
The feed plants used in the test included leaves and young anterior to the posterior end to expose the internal organs.
fruit (baby) of sweet corn (Zea mays L. (Poaceae); F1 Hybrid Furthermore, the abdomen was opened using a surgical needle
Talent, PT. Agri Makmur Pertiwi), broccoli (Brassica oleracea L. to remove the ovaries carefully. Observations were carried out
var. Italica (Brassicaceae); F1 Hybrid Broccoli Bonanza, Known- under a microscope on the number of eggs in the ovarioles.
You Seed), rice (Oryza sativa L cv. Ciherang (Gramineae)) and The data were compiled in the form of a life table. The
oil palm (Elaeis guineensis Jacq (Arecaceae)). Corn and broccoli parameters observed were as follows [16,17]:
seeds were planted using polybags with a capacity of 5 kg
containing a mixture of soil and manure (3:1). Planting of Net reproduction rate (R0)(individual/parent/generation)
(1)
rice plants starts from seeds sown on plastic trays and then, = ∑IX mx ,
after 2 weeks, transferred to plastic buckets with mixed soil
 Biology of Spodoptera frugiperda on different types of plant  3

Intrinsic growth rate (GR;) (r )(individual/parent/day) Furthermore, the nutritional indices were calculated
(2)
= (ln R0)/T , by the gravimetric method using the following formula
([18,19]):
∑XIxmx Consumption rate (CR):
Average generation period(T )(days) = , (3)
∑Ixmx
CR (g/day) = F /T , (6)
Population doubled DT(days) = ln(2)/r , (4)
Relative CR (RCR):
and
RCR (g/g body weight/day) = F /TA , (7)
Gross reproduction rate(GRR)(individual/generation)
(5) Growth Rate (GR) (g/day) = G/T,
= ∑mx ,
Relative GR (RGR):
where x: cohort age class (days); Ix: the individual prob- RGR (g/g body weight/day) = G /TA , (8)
ability of each individual at age x; mx: fecundity per indi-
vidual at age x; and IXmx: the number of offspring born in The efficiency of conversion of digested food (ECD):
the x age class.
ECD (%) = G /(F − f ) × 100%, (9)

The efficiency of conversion of ingested food (ECI):

ECI = G /F × 100%, (10)
2.4 Effect of feed plant type on food
utilization and larval growth of Approximate digestibility (AD):
S. frugiperda AD = (F − f ) /F × 100%, (11)

where G: The weight gain of larvae during the feeding

Measurement of food utilization and larval growth of S.
period, obtained from the final dry weight of the larvae
frugiperda refers to the gravimetric method [18]. Each
minus the initial dry weight; F: The amount of feed con-
treatment consisted of ten S. frugiperda instar V larvae
sumed, obtained by subtracting the initial weight from the
that had just changed cuticles. The treatment types of
final dry weight of the feed; f: dry weight of feces; T:
feed included leaves of corn, rice, broccoli, oil palm, and
feeding period; A: The average weight of larvae during
baby corn fruit as control. The experiment was arranged
treatment, obtained from the addition of the initial weight
using a Randomized Block Design with five replications.
to the final dry weight of the larvae divided by two.
The experiment was started by weighing the larvae and
feed to determine the initial wet weight, then placed indi-
vidually in a plastic cup with each type of treatment feed.
Each piece type of feed used was 4 cm × 4 cm, while corn 2.5 Data analysis
was cut with a diameter of 2 cm and a thickness of 1 cm.
The feeding periods of the larvae were 2 days and the treat- Data from biological observations were processed using
ment was ended. Furthermore, larvae, food residue, and analysis of variance, followed by Duncan’s multiple interval
feces from each plastic cup were wrapped in aluminum foil test. The life table data were processed and analyzed using
and then dried in an oven at 90°C for 48 h. Microsoft Office Excel 2010 Worksheet.
The correction factor was first calculated to obtain the
initial dry weight of the experimental larvae. Furthermore,
the correction factor was obtained by weighing the wet and
3 Results
dry weights of ten larvae. The correction factor is obtained
by calculating the dry weight of the larvae divided by the
wet weight and then multiplying by 100%. The percentage of 3.1 Effects of plant species on development
biomass content of the correction factor was then multiplied and reproduction of S. frugiperda
by the initial wet weight of the experimental larvae to
obtain the initial dry weight of the experimental larvae. The developmental phase of S. frugiperda consisted of eggs,
The initial dry weight of each feed was calculated in the larvae, pupae, and imago. The duration of the egg stage on
same way as the larvae dry weight. Finally, all values baby corn fruit, corn, rice, and broccoli leaves ranged from
were converted into dry weight values for analysis. 2.31 to 2.98 days. The duration of the egg stage laid by
4  Danar Dono et al.

imago, whose larvae were fed with rice leaves, were at from 9.26 to 10.54 days (Table 1). Furthermore, different
least 2.31 days, while corn leaves were 2.98 days longer plant species affected the pupae’s weight. The five feed
than the other four types. The type of feed used did not types tested showed significantly different values. The
affect the hatching time. Unfortunately, observation of pupae weight ranged from 0.1249 to 0.1879 g, highest in
palm leaf feed could not be conducted because only one baby corn fruit, then broccoli, baby corn leaves, and rice
abnormal pupae developed at the immature stage (Table 1). leaves, respectively (Table 2).
S. frugiperda passed six instars in the five types of feed. The sex ratio of male and female S. frugiperda that
The development length of the first to the sixth instar on emerged from the feed types of baby corn fruit was 1:1.5,
the five types of feed was significantly different. The 1.35:1) for broccoli leaf, (1:1.04) for corn leaf, and 1:1.91
shortest total duration of larval development was in the for rice leaf (Table 2). The female imago appears 2–3
baby corn fruit feed treatment (14.68 days), followed by days earlier than the male imago. Imago copulates at the
broccoli (17.60 days), corn (19.12 days), rice (20.73 days), age of 1–3 days. However, the female imago of S. frugiperda
and oil palm leaves (61.86 days) (Table 1). Meanwhile, the that emerged from the pupae in all feed treatments that did
mortality of first instar larvae was relatively high in rice not copulate for more than five days died. Hasyim et al. [20]
and oil palm leaf feed. The death of test insects between reported that in the female imago of Helicoverpa armigera,
instars I and VI were indicated with the declined amount of calling the male to copulate peaked at the age of 4 days
survival of test insect (n) in Table 1. which was thought to be related to the sexual maturity of
In oil palm leaf treatment, larvae that lived up to the female imago.
the sixth instar experienced long development and small
larval bodies. Larvae in the sixth instar that failed to enter
the prepupae stage died with symptoms of shortened and Table 2: Pupae weight and sex ratio of S. frugiperda imago
dry bodies. Oil palm leaf showed unsuitable hosts for the
development of S. frugiperda compared to the treatment of Feed type n Pupae weight Imago sex ratio
baby corn fruit, corn, rice, and broccoli leaves. Larvae that average ± SDa (g) (male:female)
feed on oil palm leaves have a long larval period indicating
Baby corn 50 0.1879 ± 0.0284d 1.0:1.50
compensation when feeding on low-quality hosts. fruit
The development of the larva ultimately affects the Corn leaf 49 0.1442 ± 0.0269b 1.0:1.04
prepupal stage and pupa formation. Moreover, the differ- Rice leaf 32 0.1249 ± 0.2020a 1.0:1.91
ence in the feed type used did not affect the duration of the Palm leaf — — —
Broccoli leaf 47 0.1656 ± 0.0273c 1.35:1.0
prepupae, which ranged from 1.78 to 2.08 days. Generally,
the duration and weight of pupae showed different values a
Numbers followed by the same letter were not significantly different
among the five feed types. The duration of pupae ranged (Duncan’s test, α = 0.05); SD – standard deviation.

Table 1: Development of S. frugiperda in several feeds

Average time required (days ± standard deviation)a

Development phase n Baby corn fruit n Corn leaf n Rice leaf n Palm leaf n Broccoli leaf

Egg b
50 2.88 ± 0.19b 50 2.98 ± 0.23b 50 2.31 ± 0.23a — — 50 2.50 ± 0.26a
Larva I 50 3.22 ± 0.42a 50 4.1 ± 0.30b 32 5.14 ± 0.91c 27 11.32 ± 1.08e 50 5.54 ± 0.50d
Larva II 50 2.1 ± 0.30a 50 2.94 ± 0.24b 32 3.18 ± 0.72b 19 9.63 ± 0.74c 50 2.14 ± 0.35b
Larva III 50 2.06 ± 0.24a 50 2.06 ± 0.24a 32 2.46 ± 0.62a 14 8.78 ± 0.84b 50 2.04 ± 0.20a
Larva IV 50 1.92 ± 0.27b 50 2.08 ± 0.27ab 32 2.58 ± 0.50b 6 8.60 ± 0.89c 50 1.72 ± 0.46a
Larva V 50 2.04 ± 0.20a 50 2.8 ± 0.45a 32 2.74 ± 0.67a 4 10.75 ± 3.30b 50 2.10 ± 0.27a
Larva VI 50 3.34 ± 0.66a 50 5.14 ± 0.76a 32 4.66 ± 1.04a 3 12.33 ± 4.62b 50 4.10 ± 0.75a
Total I–VI 50 14.68 ± 0.29a 50 19.12 ± 0.44b 32 20.76 ± 1.02b 3 61.41 ± 6.78c 50 17.64 ± 0.32ab
Prepupae 50 1.78 ± 0.42a 50 2.08 ± 0.44a 32 1.96 ± 0.93a 1 2.0 ± 0.0a 50 2.08 ± 0.35a
Pupae 50 9.26 ± 0.99a 49 9.9 ± 1.19a 32 10.54 ± 1.02a 0 — 47 10.47 ± 0.93a
a
Inline numbers followed by the same letter were not significantly different (Duncan’s test, α = 0.05); n – number of samples, bnumber of eggs
observed from adult emerge. Total I-IV represents the cumulative development time from the first instar to the sixth instar.
 Biology of Spodoptera frugiperda on different types of plant  5

The lifespan of imago ± SD (days) (n)

The number of eggs laid by each female per day in the

17.9 ± 3.8(15)b
15.3 ± 6.1(15)b

16.7 ± 5.7(15)b
baby corn treatment was higher than that in the other

9.9 ± 3.8(15)a
treatments but not significantly different from the corn

Female
leaf treatment. In the total number of eggs laid by females,

—
the baby corn treatment yields the highest number and is
different from the other treatments. However, the total

19.9 ± 2.7(15)ab

22.5 ± 4.8(15)b
22.6 ± 5.9(11)b
18.7 ± 3.3(15)a
number of eggs produced in the corn leaf, rice leaf, and
broccoli leaf treatments was not significantly different from
others. It was suggested that the preference for food does not

Male

—
significantly impact the total number of eggs produced per
female throughout their lifespan when using leaves (espe-

Description: SD – standard deviation; n – number of test insects; Numbers followed by the same letter are not significantly different (Duncan’s test, α = 0.05).
cially corn, rice, and broccoli) as food sources. Furthermore,
the insect fertility test was lowest in the rice leaf treatment,

18.4 ± 38.3a
eggs (egg)

5.7 ± 20.9a
∑ Ovarian

3.3 ± 10.2a

1.6 ± 3.4a
and the other three treatments (corn leaf, baby corn, and
broccoli leaf) were not significantly different from the other.

—
This indicated that among several test parameters (oviposi-
tion period, fecundity, fertility, and imago lifespan), the baby

Fertility (%)

86.4 ± 35.1b
95.4 ± 13.4b
54.6 ± 27.0a
98.0 ± 4.6b
corn treatment has the most significant impact compared to
the other treatments. The use of broccoli leaves is not signifi-

—
cantly different when compared to corn leaves (Table 3).
The number of eggs laid by female imago fluctuated in
each type of feed treatment. In the treatment of baby corn
∑ Eggs/female/
Table 3: Effect of several feed types on oviposition period, fecundity, fertility, and imago life span of S. frugiperda

fruit, broccoli, and rice leaves, the number of eggs laid 138.42 ± 37.6b
137.6 ± 58.0b
70.3 ± 35.2a

81.7 ± 45.4a
peaked on the first day and then decreased. Furthermore,
day (egg)

it experienced an increase again in the final stage before the

female imago died. In contrast to the corn leaf feed treat-
—

ment, eggs peaked on the seventh day after emerging from

the pupae or the fourth day after the preoviposition period.
1088.3 ± 326.0b

684.4 ± 379.3a
544.1 ± 289.7a
713.3 ± 257.9a
female (egg)

Moreover, in the final stage, it increases again before the

female imago died (Figure 1). In all treatments tested, the
∑ Eggs/

remaining eggs were found in ovary. The number of eggs

—

remaining in ovary was highest in the rice leaf treatment

with 18.4 eggs, and the lowest was 1.6 eggs in the broccoli
leaf treatment (Table 3). This indicated that Broccoli can be
∑ – The average oviposition period for female imago of S. frugiperda

the preferred choice for female imago to lay their eggs,

period (days)

similar to their original host, which is corn plant.

Oviposition

11.5 ± 3.8b
11.3 ± 5.4b
5.2 ± 2.9a

8.1 ± 4.3a
—

3.2 Effect of feed plant species on survival

and fecundity of S. frugiperda

The life table of S. frugiperda was used to determine popu-

Pre-oviposition
period (days)

lation development. The probability of individuals living at

3.5 ± 0.7ab
3.0 ± 0.6a

3.3 ± 1.0a
4.1 ± 1.4b

all stages starting from eggs, larvae, pupae, and imago (Ix)
and the fecundity of female imago (mx) was shown from
—

the survival curve. The survival curve illustrates that the

Baby corn fruit

individual’s chances of survival decreased since the age of

Broccoli leaf
Feed type

5 days after infestation (Figure 2) in the treatment of rice

Palm leaf
Corn leaf
Rice leaf

leaf feed. Meanwhile, for the treatment of corn and broc-

coli leaf feed, the chance of survival decreased at the final
6  Danar Dono et al.

350

Average amount of egg

300

produced (egg)
250
200 Baby corn fruit
150 Corn leaf
100 Rice leaf
50 Broccoli leaf
0
1 3 5 7 9 11 13 15 17 19 21 23 25 27
Egg laying time (Day)

Figure 1: Effect of different feed types on the egg-laying time and number of eggs in the female imago of S. frugiperda.

Baby corn fruit Corn leaf

1.2 200
1.2 200

Average female fecundity per day

Proportion of surviving individuals per
Proportion of surviving individuals

Average female fecundity per day

1 1
150 150
0.8 0.8
0.6 100 0.6 100
per day (lx)

0.4

(mx)
0.4

day (Ix)
(mx)

50 50
0.2 0.2
0 0 0 0
1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61 1 6 11 16 21 26 31 36 41 46 51 56 61
Age (day) Age (day)
Ix mx
Ix mx
Average female fecundity per day (mx)

Rice leaf
Proportion of surviving individual per

1.2 100
Oil palm
1 1.2
Proportion of surviving

80
individual per day (Ix)

1
0.8
day (Ix)

60 0.8
0.6
0.6
40
0.4 0.4 Ix
0.2 20 0.2
0 0 0
1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61 1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61 65
Age (day) Age (day)

Ix mx

Broccoli leaf

1.2 350
Average femlae fecundity per day
Proportion of surviving individuals

1 300
0.8 250
200
0.6
per day (Ix)

150
0.4 100
(mx)

0.2 50
0 0
1 7 13 19 25 31 37 43 49 55
Age (day)
Ix mx

Figure 2: S. frugiperda survival curve for five types of feed; No adult S. frugiperda emerged from larvae population fed on oil palm leaves and the
mortality of young larvae is high.
 Biology of Spodoptera frugiperda on different types of plant  7

Table 4: Life table of S. frugiperda on several types of feed 3.4 Effect of plant species on the nutritional
indices of S. frugiperda
Feed type Population Parameters

R0 GRR r T DT The CR, GR, and feeding efficiency of S. frugiperda larvae

instar V are shown in Table 5. The analysis of variance
Baby corn fruit 368.60 1856.41 0.16 36.29 4.26
showed that all test parameters affected the analysis of the
Corn leaf 272.66 1022.06 0.13 42.89 5.30
Rice leaf 125.90 576.16 0.10 47.02 6.74 nutritional indices of S. frugiperda larvae. The larvae CR was
Palm leaf — — — — — higher in the treatment of baby corn fruit and broccoli leaves,
Broccoli leaf 322.75 1165.92 0.14 40.37 4.84 while the lowest was in the palm and rice leaves. The
increased CR indicated that the larvae ate more parts of the
R0 – net reproduction rate (individual/parent/generation); r – intrinsic
growth rate (individual/parent/day); T – mean generation time (days); baby corn and broccoli leaves. Furthermore, the increase in
DT – time for population to double (days); GRR – gross reproduction rate the RCR on the treatment of baby corn fruit and broccoli
(individual/generation). leaves was in line with the high CR.
The value of low CR correlates with low digestibility
(AD), which also causes lower production of feces in palm
stages of development, namely, the pre-pupae and pupae
leaves. Meanwhile, high digestibility (AD) values were
stages. In the treatment of baby corn fruit feed, it did not
shown in broccoli leaf and baby corn fruit treatment. In
decrease at the early and late stages of development.
addition, larvae consuming baby corn fruit and broccoli
The average fecundity (mx) of the female imago of
leaves showed softer or watery feces, presumably some
S. frugiperda that developed from treatment of feed baby
of the water that came out with feces.
corn fruit showed high numbers at the beginning of laying
The palm leaf treatment showed the lowest GR, while the
eggs (153.56 eggs), rice leaf (82.87 eggs), and broccoli leaf (322
highest GR was in the baby corn fruit treatment. Furthermore,
eggs). Meanwhile, in the rice leaf treatment, the female imago
the GR value correlated with the RGR. The GR is affected by
laid a few eggs in the early stages of the imago, and the
the larvae CR, where the low feed consumption caused less
number increased in the middle age of the imago (576.2
feed converted into biomass, therefore a lower GR. In line with
eggs). It continued to fluctuate with increasing age of imago
Hwang et al. [21], the suitability of larval feeds containing high
and decreased before imago died in all feed treatments.
nutrients increased the GR and development period more
quickly than larvae fed with low nutrition diets.
The efficiency conversion of food ingested (ECI) was
3.3 Effect of plant species on demographic lower on palm leaf treatment, while it was the highest in
statistics of S. frugiperda the rice leaf treatment. Likewise, the efficiency conversion
of digested food (ECD) was lower on broccoli leaf treatment,
The value of the net reproduction rate (R0) was higher in while it was the highest on the corn leaf treatment. Thus,
the baby corn fruit feed treatment at 368.60, followed by high ECI and ECD values indicate that the feed effectively
broccoli, corn, and rice leaves. The value of the intrinsic GR converts ingested and digested feed into body biomass.
showing the reproductive potential (r) in the baby corn The low ECD and ECI values in the treatment of baby
fruit feed treatment was 0.16, followed by broccoli, corn, corn fruit and broccoli leaf were thought to contain sec-
and rice leaves. The smaller mean value of the generation ondary metabolites in the feed, therefore the larvae com-
period (T) indicates how faster an organism reproduces, as pensated. In line with Rahman and Rosli [22], the content of
shown by the 36.29 baby corn fruit treatment followed by polyphenols and flavonoids in baby corn fruit is higher than
broccoli, corn, and rice leaf feed treatment. The GRR of that in mature corn. Meanwhile, the high content in broccoli
S. frugiperda was 1856.41 on baby corn fruit feed, while leaf is attributed to the presence of secondary plant meta-
the GRR was 576.16 on rice leaf feed (Table 4). bolites in the form of glucosinolate compounds. Further-
The higher intrinsic GR in the baby corn fruit feed more, it is also in line with Li et al. [23] that Plutella xylostella
treatment resulted in faster development (shorter genera- treated with artificial feed containing high glucosinate was
tion time), higher endurance, and high fecundity. A high toxic to larvae and reduced the relative growth rate (RGR).
value indicates the susceptibility of the host plant to insect The inconsistent nutritional requirements across var-
food, while a low value indicates that the host plant species ious stages of larval development and the unavailability of
is slightly resistant or tolerant to pests. It was indicated various foods can lead to an increase in the amount of food
that broccoli is suitable for S. frugiperda to grow and digested (ECD) and consumed (ECI) that must be allocated
develop, as well as baby corn fruit as feed. in the metabolic process for optimal growth [24]. High ECI
8  Danar Dono et al.

All mean values ± SD. The average value in one column followed by the same letter was not significantly different (ANOVA, followed by Duncan’s test at α = 0.05). LW – larval weight, CR – consumption rate,
RCR – Relative consumption rate, GR – Growth rate, RGR – Relative growth rate, ECI – The efficiency of conversion of ingested food, ECD – The efficiency of conversion of digested food, AD – Approximate
and ECD values indicate higher efficiency of ingested and

67.637 ± 21.724ab
70.000 ± 15.353b
58.635 ± 19.072a
85.984 ± 15.744c
81.298 ± 11.361c
digested food conversion into body biomass with a high
increase in larval weight. Silva et al. [25] also stated that

AD (%)
different host plants affected the growth, weight gain, and
efficiency of digested food conversion in S. frugiperda.

Feed utilization efficiency

41.584 ± 20.928c
27.193 ± 21.908b
15.672 ± 8.499a

12.022 ± 7.941a
9.907 ± 9.621a
4 Discussion

ECD (%)
S. frugiperda was tested on several host plants, including
corn (leaf and fruit), broccoli, rice, and oil palm. The type
of host plant has a significant effect on the development,

28.568 ± 15.358c
16.436 ± 11.735b
12.353 ± 6.282b
survival, and reproduction of S. frugiperda. S. frugiperda

6.252 ± 4.164a
7.452 ± 5.729a
reared on baby corn fruit feed showed faster larval devel-

ECI (%)
opment and reproduction and high egg fertility. In con-
trast, corn leaf feed treatment showed longer larval devel-
opment time and low reproduction.
The larval development time in the treatment of broc-

RGR (g/g/day)

0.134 ± 0.088b
0.169 ± 0.096b
0.041 ± 0.027a
0.398 ± 0.176c
0.466 ± 0.111d
coli leaf was shorter than the corn and rice leaf feed treat-
ment, in line with the higher female imago reproduction
than corn leaves and rice leaves. At the beginning of the
development of S. frugiperda, larvae treated with rice leaf feed
showed high mortality, while in the oil palm leaf, the larvae
GR

did not develop as in the other four types of feed. In our study,
Table 5: Effect of several feed types on the CR, GR, and feed utilization efficiency of S. frugiperda larvae

0.0026 ± 0.002b
0.0061 ± 0.004d

0.0007 ± 0.001a
0.0041 ± 0.003c
0.007 ± 0.003d

oil palm was not a host for this insect. In Indonesia, Herlinda
GR (g/day)

et al. [26] reported that in a feeding test, S. frugiperda can eat

oil palm leaf. Unfortunately, Herlinda’s research observation
does not continue until the insect develops to imago so that it
cannot be concluded the oil palm is suitable for the growth
and development of this insect pest. Based on the literature
RCR (g/g/day)

0.622 ± 0.246a
0.785 ± 0.425a
2.329 ± 1.207b
3.035 ± 1.288c
4.331 ± 1.479c

study, Montezano et al. [10] reported that S. frugiperda is

found on oil palm, but there is no report about the insect
survival on oil palm. So, the term host plant must be defined
accurately and carefully.
CR

Differences in the length of development, survival, and

0.009 ± 0.004a
0.033 ± 0.013b

0.041 ± 0.018b
0.014 ± 0.007a
0.056 ± 0.015c

reproduction of the insect are thought to be due to sec-

CR (g/day)

ondary metabolites, plant characteristics, and poor nutri-

tional content. For example, leaves of corn plants contain
abundant benzoxazinoid as a chemical defense against
herbivores [27]. This makes the development time of larvae
on corn leaf feed longer than baby corn fruit due to the
0.0105 ± 0.0043a
0.0184 ± 0.0036b
0.0139 ± 0.0047a
0.0242 ± 0.0035c

0.0141 ± 0.0053a

presence of secondary metabolites in the leaf.

The secondary metabolite content in rice includes
oxalic acid, tricin, schaftoside, and apigenin-C-glycoside
LW (g)

compounds, which function as deterrence, antifeeding,

and are toxic to brown planthoppers [28]. The development
inhibition of S. frugiperda larvae due to disturbed feeding
Broccoli leaf

digestibility.
Feed Type

Baby corn

activity resulted in pupal weight loss and affected egg

Palm leaf
Corn leaf
Rice leaf

reproduction in rice leaf but did not directly affect the

length of larval development. This is presumably due to
 Biology of Spodoptera frugiperda on different types of plant  9

the influence of the nutritional content and secondary Furthermore, the effect of nutritional quality on several
metabolites of rice plants. feed types tested provided information that, in the absence
The high mortality in the early stages and the failure of of corn plants (whether leaves or baby corn), broccoli
larvae to reach the pre-adult stage are thought to be due to an becomes the most preferred host for S. frugiperda com-
imbalance in the composition of nutrients and secondary meta- pared to other plants tested. The research also showed
bolites found in oil palm leaf. In line with Nurhajijah [29], palm that S. frugiperda is not yet a potential pest in oil palm.
leaves contain high levels of fiber, lignin, and ash content, This research implies that S. frugiperda could become a
which have antifeedant properties for Spodoptera litura. pest on other agricultural plants in the future, especially
Differences in plant characteristics such as higher on vegetables.
water content in baby corn fruit (90.57%) than the four
types of feed used, namely, broccoli leaf (85.24%), palm Acknowledgments: The research was funded by Universitas
leaf (84.69%), corn leaf (80.75%), and rice leaf (71.32%) Padjadjaran Internal Grant Program (HIU) through the
will affect S. frugiperda. Wei et al. [30] state that the higher Research of Doctor Dissertation (Numbers: 1427/UN6.3.1/LT/
water content in a feed will positively correlate with her- 2020 and 1595/UN6.3.1/PT.00/2021) with Danar Dono as prin-
bivorous insects for host preference. cipal investigator.
It is essential to understand the nutritional status of
the different plant food types as herbivorous insect prefer- Funding information: The research was funded by Universitas
ences. The different nutritional requirements at the early Padjadjaran Internal Grant Program (Numbers: 1427/UN6.3.1/
and late stages of larval development correspond to the LT/2020 and 1595/UN6.3.1/PT.00/2021).
damage caused. This is also supported by the study of S.
frugiperda life table in our research, which shows that corn Author contributions: The authors contributed equally for
and broccoli plants were suitable for the development of S. this research work.
frugiperda. However, the high mortality of larvae in the
early stages of rice and oil palm indicates that these plants Conflicts of interest: The authors state no conflict of interest.
are unsuitable as host plants.
The suitability of host plant for larval feed which con- Data availability statement: The datasets generated and/
tains high nutrients, increased the GR and development or analyzed during the current study are available in the
period more quickly than larvae fed with low nutrition Universitas Padjadjaran repository and can be obtained
feeds [21]. Result of this experiment showed that corn from the corresponding author upon reasonable request.
and broccoli had a significant effect on the survival, devel-
opment, and reproduction of S. frugiperda. In general,
shorter development time and high reproduction repre-
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