Biology of Spodoptera frugiperda (Lepidoptera: Noctuidae) on different types of plants feeds: Potency as a pest on various agricultural plants

Danar Dono; Retno Wulansari; Yusup Hidayat; Neneng Sri Widayani

doi:10.1515/opag-2022-0254

Article Open Access

Biology of Spodoptera frugiperda (Lepidoptera: Noctuidae) on different types of plants feeds: Potency as a pest on various agricultural plants

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

Published/Copyright: February 12, 2024

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From the journal Open Agriculture Volume 9 Issue 1

Abstract

Spodoptera frugiperda is a new invasive and highly polyphagous pest that attacks corn in Indonesia. The availability of abundant plant species allows pests to switch to other host plants to maintain their population. The aim of this research is to examine the development, reproduction, nutritional indices, and life table of S. frugiperda in several plant species. The plants tested were corn, rice, broccoli, oil palm, and baby corn as controls. Ten individual insects were used and the experiments were repeated five times for each plant species. The test results show that different types of plant feed affect the development time, imago life span, fecundity, and fertility of S. frugiperda. Baby corn fruit and broccoli had higher net reproduction value (R ₀), intrinsic growth rate (r), gross reproduction rate (GRR), shorter mean generation period (T), and population doubling time (DT) than corn and rice leaves. On oil palm leaf feed, no population parameters could be determined because no larvae developed into adults and had the lowest nutritional indices parameters, so that, oil palm could not be exploited as a host plant. Also, the nutritional indices of several feed plant species tested provided information that broccoli could be the most suitable host compared to other plants tested when there was no corn in the field.

Keywords: fall armyworm; insect biology; life table; nutritional indices; host suitability

1 Introduction

Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) was the first reported pest in 1797 in Georgia, America [1]. Furthermore, this pest in 2016 was first reported in West and Central Africa [2]. Also, S. frugiperda populations were discovered in 2018 in many locations in India [3] and Southeast Asia in 2019 [4]. In early 2019, it was reported to be discovered in Indonesia [5].

The migration ability of S. frugiperda imago is high up to 1,600 km in 30 h [6]. Hence, it spread rapidly to several areas in Indonesia. For example, it was reported in West 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 corn plants from the vegetative to generative phases.

S. frugiperda is highly polyphagous, causing economic impacts on various plants [9]. Moreover, 353 different host plant species have been reported from 76 plant families based on a thorough literature review, especially the Poaceae, Asteraceae, and Fabaceae families [10], as well as the Brassicaceae family as other host [11]. The availability of abundant hosts makes S. frugiperda populations able to create new food preferences on various plants.

In line with Subramanian and Mohankumar [12], the ability of insects to survive on diverse host plants is an adaptive mechanism for survival in ecosystems. However, the availability and quality of host plants play an important role in pest population dynamics by influencing larval and imago stage performance [13]. Furthermore, according to Chapman [14], the amount and quality of food consumed by insects affects development, reproduction, and life span.

Based on the pest management perspective, the life table is essential to determine the most vulnerable pest stage for pest control [15]. Therefore, this research was conducted to examine the development, reproduction, nutritional indices, and life table of S. frugiperda in several plant species. In addition, S. frugiperda has several alternative hosts that can be utilized apart from its primary host. As an implication, we can ascertain the potential of S. frugiperda as a pest on various plants (food sources) used in this study.

2 Materials and methods

2.1 Rearing of test insect

S. frugiperda larvae were obtained from corn plantations in Jatinangor, Sumedang, West Java. Indonesia. The larvae were reared at the Pesticide and Environmental Toxicology Laboratory, Department of Plant Pests and Diseases, Faculty of Agriculture, Universitas Padjadjaran, West Java, Indonesia. Furthermore, the larvae were kept in plastic boxes measuring 34 × 28 × 7 cm. Early instar larvae kept in plastic boxes were fed with baby corn fruit (Zea mays). Before pupation, the larvae were transferred to a plastic container with a line of paper and given sawdust as the medium for pupation. Furthermore, the pupae that had been formed were transferred to the cage (measuring 44.5 × 44.5 × 49.5 cm) until they became an imago.

The imagos were fed 10% liquid honey absorbed on a lump of cotton. Afterwards, pesticide-free corn leaves were put in bottles filled with water and placed in plastic cages as a place to lay eggs. The eggs laid by imago on corn leaves were collected daily and placed in a ventilated plastic box measuring 10 × 9 × 4.5 cm, lined with paper at the bottom. Moreover, insects were maintained every day, hence, the larvae were available for testing.

2.2 Feed plant cultivation

The feed plants used in the test included leaves and young fruit (baby) of sweet corn (Zea mays L. (Poaceae); F1 Hybrid Talent, PT. Agri Makmur Pertiwi), broccoli (Brassica oleracea L. var. Italica (Brassicaceae); F1 Hybrid Broccoli Bonanza, Known-You Seed), rice (Oryza sativa L cv. Ciherang (Gramineae)) and oil palm (Elaeis guineensis Jacq (Arecaceae)). Corn and broccoli seeds were planted using polybags with a capacity of 5 kg containing a mixture of soil and manure (3:1). Planting of rice plants starts from seeds sown on plastic trays and then, after 2 weeks, transferred to plastic buckets with mixed soil conditions. Moreover, the plants were watered regularly, and then fertilization was carried out 7 days after planting with a dose of NPK (15:15:15) fertilizer of 3 g per plant for corn and broccoli plants, as well as fertilization was carried out 14 days after transplanting for rice plants.

Replanting was carried out once a week to obtain uniform and sufficient plants to feed the larvae. Plants can be used as feed after they have more than 5 leaves or are more than 2 months old. Meanwhile, oil palm leaves were obtained from young plants in Jatinangor, West Java

2.3 Effect of feed types on biology of S. frugiperda

Larvae that emerge from newly hatched eggs (<24 h) were selected to be placed in plastic cups as many as 50 larvae for each feed type. The larvae were placed separately in plastic cups (diameter of 2.5 cm, height of 4 cm) with leaves of corn, rice, broccoli, oil palm, and baby corn fruit as control. Feed leaves and baby corn fruits were replaced daily with fresh ones.

Observations were made every day to determine the mortality and development time of larvae. In addition, the length of larval development was observed by recording the time required for S. frugiperda larvae to develop from a certain instar to the next, marked by the molting of the larval cuticle. After the larvae became pupae, observations were made, including the development time and weight of pupae, normal and abnormal conditions, as well as mortality. Weight of pupae was measured on the third day after pupation using analytical balance.

The imago that emerged from pupae were paired in a cage (diameter of 13.5 cm and height of 13 cm) where a 10% honey solution was absorbed into the cotton as food of imago. Furthermore, the corn leaves, rice leaves, and broccoli leaves 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 were dissected to reveal ovaries. A lateral incision was made in the abdomen following the midline of the thorax from the anterior to the posterior end to expose the internal organs. Furthermore, the abdomen was opened using a surgical needle to remove the ovaries carefully. Observations were carried out under a microscope on the number of eggs in the ovarioles.

The data were compiled in the form of a life table. The parameters observed were as follows [16,17]:

(1) Net reproduction rate ( R 0 ) ( individual/parent/generation ) = ∑ I X m x ,

(2) Intrinsic growth rate (GR;) ( r ) ( individual/parent/day ) = ( ln R 0 ) / T ,

(3) Average generation period ( T ) ( days ) = ∑ X I x m x ∑ I x m x ,

(4) Population doubled DT ( days ) = ln ( 2 ) / r ,

and

(5) Gross reproduction rate ( GRR ) ( individual/generation ) = ∑ m x ,

where x: cohort age class (days); I _x: the individual probability of each individual at age x; m _x: fecundity per individual at age x; and I _X m _x: the number of offspring born in the x age class.

2.4 Effect of feed plant type on food utilization and larval growth of S. frugiperda

Measurement of food utilization and larval growth of S. frugiperda refers to the gravimetric method [18]. Each treatment consisted of ten S. frugiperda instar V larvae that had just changed cuticles. The treatment types of feed included leaves of corn, rice, broccoli, oil palm, and baby corn fruit as control. The experiment was arranged using a Randomized Block Design with five replications. The experiment was started by weighing the larvae and feed to determine the initial wet weight, then placed individually in a plastic cup with each type of treatment feed. Each piece type of feed used was 4 cm × 4 cm, while corn 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 treatment was ended. Furthermore, larvae, food residue, and feces from each plastic cup were wrapped in aluminum foil and then dried in an oven at 90°C for 48 h.

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 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 biomass content of the correction factor was then multiplied by the initial wet weight of the experimental larvae to obtain the initial dry weight of the experimental larvae. The initial dry weight of each feed was calculated in the same way as the larvae dry weight. Finally, all values were converted into dry weight values for analysis.

Furthermore, the nutritional indices were calculated by the gravimetric method using the following formula ([18,19]):

Consumption rate (CR):

(6) CR ( g / day ) = F / T ,

Relative CR (RCR):

(7) RCR ( g / g body weight / day ) = F / TA ,

Growth Rate (GR) (g/day) = G/T,

Relative GR (RGR):

(8) RGR ( g / g body weight / day ) = G / TA ,

The efficiency of conversion of digested food (ECD):

(9) ECD ( % ) = G / ( F − f ) × 100 % ,

The efficiency of conversion of ingested food (ECI):

(10) ECI = G / F × 100 % ,

Approximate digestibility (AD):

(11) AD = ( F − f ) / F × 100 % ,

where G: The weight gain of larvae during the feeding period, obtained from the final dry weight of the larvae minus the initial dry weight; F: The amount of feed consumed, obtained by subtracting the initial weight from the final dry weight of the feed; f: dry weight of feces; T: feeding period; A: The average weight of larvae during treatment, obtained from the addition of the initial weight to the final dry weight of the larvae divided by two.

2.5 Data analysis

Data from biological observations were processed using analysis of variance, followed by Duncan’s multiple interval test. The life table data were processed and analyzed using Microsoft Office Excel 2010 Worksheet.

3 Results

3.1 Effects of plant species on development and reproduction of S. frugiperda

The developmental phase of S. frugiperda consisted of eggs, larvae, pupae, and imago. The duration of the egg stage on baby corn fruit, corn, rice, and broccoli leaves ranged from 2.31 to 2.98 days. The duration of the egg stage laid by imago, whose larvae were fed with rice leaves, were at least 2.31 days, while corn leaves were 2.98 days longer than the other four types. The type of feed used did not affect the hatching time. Unfortunately, observation of palm leaf feed could not be conducted because only one abnormal pupae developed at the immature stage (Table 1).

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

^aInline 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.

S. frugiperda passed six instars in the five types of feed. The development length of the first to the sixth instar on the five types of feed was significantly different. The shortest total duration of larval development was in the baby corn fruit feed treatment (14.68 days), followed by broccoli (17.60 days), corn (19.12 days), rice (20.73 days), and oil palm leaves (61.86 days) (Table 1). Meanwhile, the mortality of first instar larvae was relatively high in rice and oil palm leaf feed. The death of test insects between instars I and VI were indicated with the declined amount of survival of test insect (n) in Table 1.

In oil palm leaf treatment, larvae that lived up to 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 dry bodies. Oil palm leaf showed unsuitable hosts for the development of S. frugiperda compared to the treatment of baby corn fruit, corn, rice, and broccoli leaves. Larvae that feed on oil palm leaves have a long larval period indicating compensation when feeding on low-quality hosts.

The development of the larva ultimately affects the prepupal stage and pupa formation. Moreover, the difference in the feed type used did not affect the duration of the prepupae, which ranged from 1.78 to 2.08 days. Generally, the duration and weight of pupae showed different values among the five feed types. The duration of pupae ranged from 9.26 to 10.54 days (Table 1). Furthermore, different plant species affected the pupae’s weight. The five feed types tested showed significantly different values. The pupae weight ranged from 0.1249 to 0.1879 g, highest in baby corn fruit, then broccoli, baby corn leaves, and rice leaves, respectively (Table 2).

Table 2

Pupae weight and sex ratio of S. frugiperda imago

Feed type	n	Pupae weight average ± SD^a (g)	Imago sex ratio (male:female)
Baby corn fruit	50	0.1879 ± 0.0284d	1.0:1.50
Corn leaf	49	0.1442 ± 0.0269b	1.0:1.04
Rice leaf	32	0.1249 ± 0.2020a	1.0:1.91
Palm leaf	—	—	—
Broccoli leaf	47	0.1656 ± 0.0273c	1.35:1.0

^aNumbers followed by the same letter were not significantly different (Duncan’s test, α = 0.05); SD – standard deviation.

The sex ratio of male and female S. frugiperda that emerged from the feed types of baby corn fruit was 1:1.5, 1.35:1) for broccoli leaf, (1:1.04) for corn leaf, and 1:1.91 for rice leaf (Table 2). The female imago appears 2–3 days earlier than the male imago. Imago copulates at the age of 1–3 days. However, the female imago of S. frugiperda that emerged from the pupae in all feed treatments that did not copulate for more than five days died. Hasyim et al. [20] reported that in the female imago of Helicoverpa armigera, calling the male to copulate peaked at the age of 4 days which was thought to be related to the sexual maturity of the female imago.

The number of eggs laid by each female per day in the baby corn treatment was higher than that in the other treatments but not significantly different from the corn 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 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 significantly impact the total number of eggs produced per female throughout their lifespan when using leaves (especially corn, rice, and broccoli) as food sources. Furthermore, the insect fertility test was lowest in the rice leaf treatment, 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 (oviposition period, fecundity, fertility, and imago lifespan), the baby corn treatment has the most significant impact compared to the other treatments. The use of broccoli leaves is not significantly different when compared to corn leaves (Table 3).

Table 3

Effect of several feed types on oviposition period, fecundity, fertility, and imago life span of S. frugiperda

Feed type	Pre-oviposition period (days)	Oviposition period (days)	∑ Eggs/female (egg)	∑ Eggs/female/day (egg)	Fertility (%)	∑ Ovarian eggs (egg)	The lifespan of imago ± SD (days) (n)
Feed type	Pre-oviposition period (days)	Oviposition period (days)	∑ Eggs/female (egg)	∑ Eggs/female/day (egg)	Fertility (%)	∑ Ovarian eggs (egg)	Male	Female
Baby corn fruit	3.0 ± 0.6a	11.3 ± 5.4b	1088.3 ± 326.0b	138.42 ± 37.6b	98.0 ± 4.6b	5.7 ± 20.9a	19.9 ± 2.7(15)ab	15.3 ± 6.1(15)b
Corn leaf	3.5 ± 0.7ab	5.2 ± 2.9a	544.1 ± 289.7a	137.6 ± 58.0b	95.4 ± 13.4b	3.3 ± 10.2a	18.7 ± 3.3(15)a	9.9 ± 3.8(15)a
Rice leaf	4.1 ± 1.4b	11.5 ± 3.8b	713.3 ± 257.9a	70.3 ± 35.2a	54.6 ± 27.0a	18.4 ± 38.3a	22.6 ± 5.9(11)b	17.9 ± 3.8(15)b
Palm leaf	—	—	—	—	—	—	—	—
Broccoli leaf	3.3 ± 1.0a	8.1 ± 4.3a	684.4 ± 379.3a	81.7 ± 45.4a	86.4 ± 35.1b	1.6 ± 3.4a	22.5 ± 4.8(15)b	16.7 ± 5.7(15)b

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).

∑ – The average oviposition period for female imago of S. frugiperda

The number of eggs laid by female imago fluctuated in each type of feed treatment. In the treatment of baby corn fruit, broccoli, and rice leaves, the number of eggs laid peaked on the first day and then decreased. Furthermore, it experienced an increase again in the final stage before the female imago died. In contrast to the corn leaf feed treatment, eggs peaked on the seventh day after emerging from the pupae or the fourth day after the preoviposition period. Moreover, in the final stage, it increases again before the female imago died (Figure 1). In all treatments tested, the 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 preferred choice for female imago to lay their eggs, similar to their original host, which is corn plant.

Figure 1

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

3.2 Effect of feed plant species on survival and fecundity of S. frugiperda

The life table of S. frugiperda was used to determine population development. The probability of individuals living at 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 individual’s chances of survival decreased since the age of 5 days after infestation (Figure 2) in the treatment of rice leaf feed. Meanwhile, for the treatment of corn and broccoli leaf feed, the chance of survival decreased at the final stages of development, namely, the pre-pupae and pupae stages. In the treatment of baby corn fruit feed, it did not decrease at the early and late stages of development.

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.

The average fecundity (mx) of the female imago of S. frugiperda that developed from treatment of feed baby corn fruit showed high numbers at the beginning of laying eggs (153.56 eggs), rice leaf (82.87 eggs), and broccoli leaf (322 eggs). Meanwhile, in the rice leaf treatment, the female imago laid a few eggs in the early stages of the imago, and the number increased in the middle age of the imago (576.2 eggs). It continued to fluctuate with increasing age of imago and decreased before imago died in all feed treatments.

3.3 Effect of plant species on demographic statistics of S. frugiperda

The value of the net reproduction rate (R ₀) was higher in the baby corn fruit feed treatment at 368.60, followed by broccoli, corn, and rice leaves. The value of the intrinsic GR showing the reproductive potential (r) in the baby corn fruit feed treatment was 0.16, followed by broccoli, corn, and rice leaves. The smaller mean value of the generation period (T) indicates how faster an organism reproduces, as shown by the 36.29 baby corn fruit treatment followed by broccoli, corn, and rice leaf feed treatment. The GRR of S. frugiperda was 1856.41 on baby corn fruit feed, while the GRR was 576.16 on rice leaf feed (Table 4).

Table 4

Life table of S. frugiperda on several types of feed

Feed type	Population Parameters
Feed type	R ₀	GRR	r	T	DT
Baby corn fruit	368.60	1856.41	0.16	36.29	4.26
Corn leaf	272.66	1022.06	0.13	42.89	5.30
Rice leaf	125.90	576.16	0.10	47.02	6.74
Palm leaf	—	—	—	—	—
Broccoli leaf	322.75	1165.92	0.14	40.37	4.84

R ₀ – net reproduction rate (individual/parent/generation); r – intrinsic growth rate (individual/parent/day); T – mean generation time (days); DT – time for population to double (days); GRR – gross reproduction rate (individual/generation).

The higher intrinsic GR in the baby corn fruit feed treatment resulted in faster development (shorter generation time), higher endurance, and high fecundity. A high value indicates the susceptibility of the host plant to insect food, while a low value indicates that the host plant species is slightly resistant or tolerant to pests. It was indicated that broccoli is suitable for S. frugiperda to grow and develop, as well as baby corn fruit as feed.

3.4 Effect of plant species on the nutritional indices of S. frugiperda

The CR, GR, and feeding efficiency of S. frugiperda larvae instar V are shown in Table 5. The analysis of variance showed that all test parameters affected the analysis of the nutritional indices of S. frugiperda larvae. The larvae CR was higher in the treatment of baby corn fruit and broccoli leaves, while the lowest was in the palm and rice leaves. The increased CR indicated that the larvae ate more parts of the baby corn and broccoli leaves. Furthermore, the increase in the RCR on the treatment of baby corn fruit and broccoli leaves was in line with the high CR.

Table 5

Effect of several feed types on the CR, GR, and feed utilization efficiency of S. frugiperda larvae

Feed Type		CR		GR		Feed utilization efficiency
Feed Type	LW (g)	CR (g/day)	RCR (g/g/day)	GR (g/day)	RGR (g/g/day)	ECI (%)	ECD (%)	AD (%)
Baby corn	0.0242 ± 0.0035c	0.056 ± 0.015c	4.331 ± 1.479c	0.007 ± 0.003d	0.466 ± 0.111d	12.353 ± 6.282b	15.672 ± 8.499a	81.298 ± 11.361c
Corn leaf	0.0139 ± 0.0047a	0.033 ± 0.013b	3.035 ± 1.288c	0.0061 ± 0.004d	0.398 ± 0.176c	16.436 ± 11.735b	27.193 ± 21.908b	67.637 ± 21.724ab
Rice leaf	0.0141 ± 0.0053a	0.014 ± 0.007a	0.622 ± 0.246a	0.0041 ± 0.003c	0.169 ± 0.096b	28.568 ± 15.358c	41.584 ± 20.928c	70.000 ± 15.353b
Palm leaf	0.0105 ± 0.0043a	0.009 ± 0.004a	0.785 ± 0.425a	0.0007 ± 0.001a	0.041 ± 0.027a	6.252 ± 4.164a	12.022 ± 7.941a	58.635 ± 19.072a
Broccoli leaf	0.0184 ± 0.0036b	0.041 ± 0.018b	2.329 ± 1.207b	0.0026 ± 0.002b	0.134 ± 0.088b	7.452 ± 5.729a	9.907 ± 9.621a	85.984 ± 15.744c

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 digestibility.

The value of low CR correlates with low digestibility (AD), which also causes lower production of feces in palm leaves. Meanwhile, high digestibility (AD) values were shown in broccoli leaf and baby corn fruit treatment. In addition, larvae consuming baby corn fruit and broccoli leaves showed softer or watery feces, presumably some of the water that came out with feces.

The palm leaf treatment showed the lowest GR, while the highest GR was in the baby corn fruit treatment. Furthermore, the GR value correlated with the RGR. The GR is affected by the larvae CR, where the low feed consumption caused less feed converted into biomass, therefore a lower GR. In line with Hwang et al. [21], the suitability of larval feeds containing high nutrients increased the GR and development period more quickly than larvae fed with low nutrition diets.

The efficiency conversion of food ingested (ECI) was lower on palm leaf treatment, while it was the highest in the rice leaf treatment. Likewise, the efficiency conversion of digested food (ECD) was lower on broccoli leaf treatment, while it was the highest on the corn leaf treatment. Thus, high ECI and ECD values indicate that the feed effectively converts ingested and digested feed into body biomass.

The low ECD and ECI values in the treatment of baby corn fruit and broccoli leaf were thought to contain secondary metabolites in the feed, therefore the larvae compensated. In line with Rahman and Rosli [22], the content of polyphenols and flavonoids in baby corn fruit is higher than that in mature corn. Meanwhile, the high content in broccoli leaf is attributed to the presence of secondary plant metabolites in the form of glucosinolate compounds. Furthermore, it is also in line with Li et al. [23] that Plutella xylostella treated with artificial feed containing high glucosinate was toxic to larvae and reduced the relative growth rate (RGR).

The inconsistent nutritional requirements across various stages of larval development and the unavailability of various foods can lead to an increase in the amount of food digested (ECD) and consumed (ECI) that must be allocated in the metabolic process for optimal growth [24]. High ECI and ECD values indicate higher efficiency of ingested and digested food conversion into body biomass with a high increase in larval weight. Silva et al. [25] also stated that different host plants affected the growth, weight gain, and efficiency of digested food conversion in S. frugiperda.

4 Discussion

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, survival, and reproduction of S. frugiperda. S. frugiperda reared on baby corn fruit feed showed faster larval development and reproduction and high egg fertility. In contrast, corn leaf feed treatment showed longer larval development time and low reproduction.

The larval development time in the treatment of broccoli leaf was shorter than the corn and rice leaf feed treatment, 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 did not develop as in the other four types of feed. In our study, oil palm was not a host for this insect. In Indonesia, Herlinda 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 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.

Differences in the length of development, survival, and reproduction of the insect are thought to be due to secondary metabolites, plant characteristics, and poor nutritional 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 presence of secondary metabolites in the leaf.

The secondary metabolite content in rice includes oxalic acid, tricin, schaftoside, and apigenin-C-glycoside compounds, which function as deterrence, antifeeding, and are toxic to brown planthoppers [28]. The development inhibition of S. frugiperda larvae due to disturbed feeding activity resulted in pupal weight loss and affected egg reproduction in rice leaf but did not directly affect the length of larval development. This is presumably due to the influence of the nutritional content and secondary metabolites of rice plants.

The high mortality in the early stages and the failure of larvae to reach the pre-adult stage are thought to be due to an imbalance in the composition of nutrients and secondary metabolites found in oil palm leaf. In line with Nurhajijah [29], palm leaves contain high levels of fiber, lignin, and ash content, which have antifeedant properties for Spodoptera litura.

Differences in plant characteristics such as higher water content in baby corn fruit (90.57%) than the four types of feed used, namely, broccoli leaf (85.24%), palm leaf (84.69%), corn leaf (80.75%), and rice leaf (71.32%) will affect S. frugiperda. Wei et al. [30] state that the higher water content in a feed will positively correlate with herbivorous insects for host preference.

It is essential to understand the nutritional status of the different plant food types as herbivorous insect preferences. The different nutritional requirements at the early and late stages of larval development correspond to the damage caused. This is also supported by the study of S. frugiperda life table in our research, which shows that corn and broccoli plants were suitable for the development of S. frugiperda. However, the high mortality of larvae in the early stages of rice and oil palm indicates that these plants are unsuitable as host plants.

The suitability of host plant for larval feed which contains high nutrients, increased the GR and development period more quickly than larvae fed with low nutrition feeds [21]. Result of this experiment showed that corn and broccoli had a significant effect on the survival, development, and reproduction of S. frugiperda. In general, shorter development time and high reproduction represent higher suitability of host plant. The test results showed that broccoli plants were first reported to be suitable for the survival of S. frugiperda. So far, the Broccoli plant (Brassica oleracea L. var. Italica) has never been reported as a host for this insect. As an implication, S. frugiperda has the potential to become a pest on other vegetable plants. Result of this research in line with Wang et al. [31], indicate that older larvae are very voracious and can cause serious losses to Chinese Cabbage (Brassica pekinensis (Lour.) Rupr) var. Qinza 2 (Brassicaceae), although younger larvae experience high mortality and only produce 5.3% imago with a sex ratio of 2:1 (male:female).

5 Conclusion

Different types of feed affect the developmental time, imago life span, fecundity, and fertility of S. frugiperda. Furthermore, the effect of nutritional quality on several feed types tested provided information that, in the absence of corn plants (whether leaves or baby corn), broccoli becomes the most preferred host for S. frugiperda compared to other plants tested. The research also showed that S. frugiperda is not yet a potential pest in oil palm. This research implies that S. frugiperda could become a pest on other agricultural plants in the future, especially on vegetables.

Acknowledgments

The research was funded by Universitas Padjadjaran Internal Grant Program (HIU) through the Research of Doctor Dissertation (Numbers: 1427/UN6.3.1/LT/2020 and 1595/UN6.3.1/PT.00/2021) with Danar Dono as principal investigator.

Funding information: The research was funded by Universitas Padjadjaran Internal Grant Program (Numbers: 1427/UN6.3.1/LT/2020 and 1595/UN6.3.1/PT.00/2021).
Author contributions: The authors contributed equally for this research work.
Conflicts of interest: The authors state no conflict of interest.
Data availability statement: The datasets generated and/or analyzed during the current study are available in the Universitas Padjadjaran repository and can be obtained from the corresponding author upon reasonable request.

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Received: 2023-07-08

Revised: 2023-11-19

Accepted: 2024-01-03

Published Online: 2024-02-12

This work is licensed under the Creative Commons Attribution 4.0 International License.

Articles in the same Issue

https://doi.org/10.1515/opag-2022-0254

Keywords for this article

fall armyworm; insect biology; life table; nutritional indices; host suitability

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