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Management of Spodoptera frugiperda: History
Please note this is an old version of this entry, which may differ significantly from the current revision.
Subjects: Entomology

The fall armyworm of maize, Spodoptera frugiperda (J. E. Smith) (Lepidoptera; Noctuidae) is capable of causing a 100% yield loss due to its unforeseen occurrence from the seedling to the cob formation stage. To manage this serious pest, maize growers are tending to apply a high dosage of pesticides. This indiscriminate usage of pesticides has resulted in an unacceptable amount of insect resurgence in maize, harming maize production and consumption.

  • agronomic practices
  • crop damage
  • integrated pest management
  • maize

1. Agronomic Management

1.1. Sowing Window

Planting dates alter the growing environment of crop plants [27]. Pests are an integral part of the agro-ecosystem. Weather variables, such as temperature, relative humidity, rainfall, etc., have a significant impact on insect infestation and population growth. Insect pests’ survival, development, and reproductive capacity are all influenced by environmental variables [28]. Usually, fall armyworm appears late in the growing season. Hence, to avoid its infestation, late sowing in kharif crops should be avoided. Furthermore, seeding kharif maize should be limited to the earliest and narrowest sowing windows to disrupt the host range’s continuous availability because early harvest helps maize ears avoid the increased armyworm concentrations that emerge later in the season [29]. Furthermore, the study conducted by Teare et al. [30] also confirms the advantages of early sowing by stating that fall armyworm of maize was not a severe problem in early-planted maize. Kandel and Poudel [31] stated that fall armyworm is likely to affect the adoption of late-maturing hybrids and late-planted maize crops in Nepal. A recent study by Food and Agriculture Organizations (2018) [32] found considerable yield losses in maize due to the occurrence of fall armyworms in late-planted maize plots compared to nearby plots that were planted earlier. Therefore, early planting and adoption of short-duration genotypes are precise practices in endemic patches of fall armyworm in US corn belts [33]. According to Bhusal and Chapagain [34], planting maize earlier than the actual date will avoid the pest’s arrival time, allowing the crop to be protected or less afflicted. Based on the limited published work with regards to the influence of sowing windows on the pest reaction of fall armyworm in maize, it can be concluded that early sowing denies the entry and establishment of fall armyworm in the maize ecosystem so that possible yield loss can be avoided.

1.2. Tillage and Land Preparation

Tillage and soil management have a significant impact on the agro–pest ecosystem’s dynamics [35]. It is a fact that continuous and intensive land disturbance has deleterious impacts on soil by favouring erosion, organic matter depletion, and negative impacts on overall soil organisms [36]. Despite the fact that tillage has a number of negative effects on agroecosystem sustainability, it is the most widely used agro-technique in crop production worldwide to prepare fields for planting and manage pests [37]. This is especially true in the case of the management of the fall armyworm in maize. The grown larvae fall to the ground to pupate, making a reddish-brown oval cocoon that is about 2–3 cm long. In light soil, they can burrow 2–8 cm, whereas in hard soil, they can spin a webbed cocoon under leaf detritus. These pest propagules in turn contribute to life cycle completion and further multiplication. 
Based on the findings of other studies, tillage may be a viable fall armyworm control strategy with minimal soil disturbance and residue retention that can be easily integrated with existing fall armyworm control efforts while also promoting sustainable intensification and climate change adaptation through conservation agriculture [42,43]. Furthermore, Harrison et al. [44] also reviewed that agroecological approaches to pest management, such as land tilling, bund planting, and soil fertility management, offer appropriate and low-cost pest control strategies that can be readily integrated into existing efforts of pest management strategies.

1.3. Nutrient Management

Balanced crop nutrition is a key factor in producing healthy plants and crops, and a healthy plant and vigorous crop growth usually resist biotic and abiotic stresses [27,44]. However, imbalanced crop nutrition paves the way for increased pest damage [45]. Morales et al. [46] reported that the introduction of inorganic fertilisers encouraged herbivorous pests in maize fields, including fall armyworm. A study conducted by Morales et al. [46] confirmed that crops fertilised with inorganic fertilizers had a higher incidence of aphids than organically fertilized crops. Inorganic sources of plant nutrients, such as chemical fertilizer amendments, assure the instantaneous supply of plant nutrients immediately after their application, especially with nitrogen. This spike in nitrogen content in the plant tissues results in attractiveness to insect herbivores and was evident from several studies conducted [45,46].  Therefore, it is imperative to amend the soil with organic manures, crop rotation with restorative field crops, green manuring, and inclusion of bio-inoculate-mediated crop nutrient modules to efficiently manage the fall armyworm.

1.4. Cropping System Measures

Growing crops in combination or succession is a vague practice in modern intensive agriculture. Cultivation of different crop species on the same piece of land imparts ecological diversity and prosperity. Bi-cropping is the practice of growing two crops with dissimilar growth habits on the same piece of land in a given period [49]. In addition to its numerous advantages over solitary cropping, bi-cropping practices in the traditional farming system give insurance against pests and unusual weather [48]. By increasing soil quality, stimulating vigorous plant development [50], restricting insect migration [51], obstructing egg-laying through visual or chemical disruption [52], and providing habitat for natural enemies, bi-cropping can lessen pest damage [53].  In comparison to mono-cropped maize, intercropping maize with leguminous crops resulted in a considerable reduction in fall armyworm, notably during the early growth stages of the maize up to tasselling. Hence, in devising integrated management strategies against fall armyworm utilisation of plant species that produce semiochemicals may be incorporated into the cropping system. Furthermore, numerous research studies from around the world have found that leguminous intercrops are beneficial in controlling fall armyworm infestations in maize [53,57].

1.5. Trap Cropping

Trap crops are designed to attract pests away from cash crops, protecting them from assault [58]. The trap crop can be from the same family group as the main crop or from a separate one. Trap crops can be planted in two ways: perimeter trap cropping and row intercropping. The planting of trap crops around the perimeter of the primary cash crop is known as perimeter trap cropping (border trap cropping). It guards against pest attacks from all sides of the field. It is most effective against pests that are found along the farm’s perimeter. The trap crop is planted in alternating rows with the main crop in intercropped rows. Trap cropping has several pest management advantages. Because harm to main crops is reduced when trap crops effectively attract pest populations, major crops rarely require insecticide treatments [58]. Insect pests can be treated in a confined region rather than the entire field when they are highly concentrated in trap crops. Savings from reduced pest attack and insecticide use may far surpass the cost of sustaining non-profitable crops. The fall armyworm is a relatively recent pest in most of the world’s maize-growing regions. Information on trap cropping practice for the management of fall armyworm is scarce. On the other hand, Mooventhan et al. [59] advocate sowing 3–4 rows of Napier grass surrounding maize fields and spraying with 5% neem seed kernel extract or azadirachtin 1500 ppm as soon as the trap crop displays symptoms of fall armyworm damage. Furthermore, Gueraet al. [60] noticed more oviposited eggs on Brachiaria hybrid cv. Mulato II (Family: Poaceae), Panicum maximum cv. Mombasa (Family: Poaceae), and Panicum maximum cv. Tanzania (Family: Poaceae) than on maize. Trap crops could also be used in the “Push–Pull” cropping method, which involves intercropping pest-repellent (“push”) plant species (e.g., Desmodium spp.) with a pest-attractive trap (“pull”) plant species on the borders (e.g., napier grass (Pennisetum purpureum Schumach.) or Brachiaria spp.). Farmers in East Africa who used the Push–Pull technique completely saw an 86% reduction in fall armyworm infestation and crop loss as well as a 2.7-fold increase in production [53].

1.6. Pheromones Traps

Pheromones are scents produced by males or females that activate one or more behavioural reactions in the opposite sex, attracting males and females to mate. The use of pheromones primarily reduces insect populations in the given locality by way of lowering their reproduction rate because these chemicals confuse insects and get them trapped [61]. The female pheromone of S. frugiperda which attracts male moths consists of the major component (Z)-9-tetradecenyl acetate (Z9–14:Ac) and the minor component (Z)-7-dodecenyl acetate (Z7–12:Ac) [62]. The minor component (E)-7-dodecenyl acetate (E7–12:Ac) shows geographic variation and has so far only been found in females from Brazil. Regarding pest management, it is vital to remember that fall armyworm is made up of two strains (corn and rice) that have different pheromones [13]. In the case of fall armyworm, the female’s sex pheromone is commercially available in numerous countries [63], and pheromones have long been used to monitor the male population [64]. Monitoring with pheromone traps is useful because pest infestation varies from farm to farm and over time. Knowing when and where the adult pest is active and plentiful gives an early warning system that allows for field sampling and treatment. The grower was able to prevent wasteful pesticide applications or time-consuming samples by learning about the presence or absence of pests, and he was also given the warning to safeguard crops when moth flight was first identified [63]. Adult male moths are caught in traps, but larvae cause plant harm. As a result, we cannot simply count the number of moths in the traps while ignoring other elements, such as temperature, crop stage, and even natural control. Wind speed and temperature are favourably connected to trap captures, but relative humidity is adversely correlated. Monitoring fall armyworm adults with pheromone traps is the most effective way to determine the number of pesticide applications required to manage the pest in maize [63].

2. Biotechnological Approach

2.1. Plant Resistance

Host plant resistance is a low-cost and potentially effective method of insect pest control. It is generally inexpensive, durable, non-polluting, and locally adaptable, which helps in sustainable production. Fall-armyworm-resistant genotypes have been developed using a variety of plant morphological traits that contribute to antixenosis. Among them, a classic study by Sanches et al. [65] found the larval phase, metabolised food, and insect stool bulk as critical factors that contribute to fall armyworm resistance. From their study, maize genotypes BOZM 260, PA 091, and PARA 172 have emerged as promising sources of resistance to the fall armyworm. Similarly, Chen et al. [66] stated that maize accession Mp708 and FAW7050 were resistant to fall armyworm due to enhanced defence protein, greater amino acid and glucose content, and constitutive jasmonic acid accumulation. A study by Smith et al. [61] indicated that (E)-β-caryophyllene, a terpenoid released constitutively in a maize line, Mp708 is responsible for the demonstration of fall armyworm resistance. Furthermore, a study conducted by Ni et al. [67] also identified maize germplasm Mp708 and FAW7061 as highly resistant accession to fall armyworm infestation. Many fall-armyworm-resistant sources have been identified by researchers all over the world; these resistance sources can be used as possible parents in the development of resistant/tolerant varieties and hybrids in the future breeding programme to reduce seedling harm.

2.2. Genetic Engineering

Employing synthetic chemical pesticides is a commonly observed phenomenon in normal cultivation practices. In chemical control, pests come into contact with the lethal substance and are killed. The chemicals are applied only when characteristic symptoms are noticed on the plant. This technique is not normally effective for fall armyworm management because caterpillars are usually found in corn whorls, where they are generally protected from insecticide treatments [71]. The fall armyworm, on the other hand, is resistant to over 30 active components of insecticides from all main classes on a global scale. Therefore, the utilization of genetically engineered maize that resists fall armyworm is a feasible method of pest management [72]. The efficacy of using genetically engineered crops in managing the fall armyworm was reported in China [73]. To control caterpillar pests, genetically engineered crop types expressing insecticidal crystalline (Cry) or vegetative insecticidal proteins (Vip) generated from Bacillus thuringiensis (Bt) that are selectively poisonous to distinct insect species are planted in many regions of the world, with adoption rates of over 80% [74]. Mass adoption, together with high control efficacy provided by genetic engineering, has led to area-wide population reductions in some pests that were already established [72]. Furthermore, the use of Bt corn hybrids is widely used to manage fall armyworm in America [71,72]. Growing Bt crops has helped to minimize pollution since they can reduce the use of chemical insecticides and assist natural pest management due to the narrow spectrum of activity of the deployed Cry and Vip proteins [74]. Genetically modified maize, according to Li et al. [73], should not be viewed as a single instrument for managing fall armyworm, but rather as a complement to other measures, such as biological control, cultural controls, and judicious use of chemical insecticides. Studies in this line have been meagre until now. This might be the one potential option for the management of fall armyworm which could be incorporated by taking into consideration only the biosafety and policy issues of respective maize-growing regions. A good insect resistance management plan, particularly for the fall armyworm, is critical and can be achieved through gene pyramiding approaches [73,75].

2.3. Gene Editing Approach (CRISPR-Cas System)

Even though transgenic (Bt) crops have provided significant crop protection benefits, the technology has been plagued by insect resistance, leading to the development of newer biotechnological approaches to insect pest management, such as gene editing (RNA interference (RNAi); gene drives, and, most recently, the CRISPR-Cas9 system) [76,77]. Gene editing technology can be used to validate gene activity, allowing researchers to better understand the resistance mechanism and design new pest management strategies. In 2013, CRISPR-Cas application in plants was successfully achieved in the lab [78].
Wu [79] revealed CRISPR/Cas9-mediated deletion of the abdominal-A homeotic gene in the fall armyworm, implying that the CRISPR/Cas9 technology is highly efficient in editing the fall armyworm genome based on his findings. For genome editing in the fall armyworm Spodoptera frugiperda, CRISPR/Cas9-mediated site-specific mutagenesis of three target genes—two marker genes (Biogenesis of lysosome-related organelles complex 1 subunit 2(BLOS2) and tryptophan 2, 3-dioxygenase(TO)), and a developmental gene, E93 (a key ecdysone-induced transcription factor that promotes adult development)—was performed. These mutational studies demonstrated there is a need to improve genome editing in lepidopteran and other non-model insects by using different approaches [80].

3. Chemical Management

Poison Baits

Poisonous baits are pesticides that can be mixed with some nontoxic food additives and have a lethal effect on targeted pests. In fall armyworm management, the application of poison bait in the whorls at vegetative and broadcasting in the mature crop has also shown a good response to controlling infestation [33]. In India, Patil et al. [58] elaborated on the protocol for the preparation of poison bait, wherein 5.0 kg of jaggery was mixed with 4–5 L of water. To this solution, 625.0 mL of monocrotophos 36 SL was added. This solution was further mixed with 50 kg of rice or wheat bran and packed in gunny or plastic bags and allowed to ferment for 48 h. Application of this fermented bait either through broadcasting or placing in maize whorls, preferably in the evening hours, significantly reduced fall armyworm incidence in maize.

Insecticides

The application of varieties of chemicals in the management of crop pests is an ambiguous practice [81,82,83]. Based on the mode of action, insecticides are divided into systemic and contact insecticides. The fall armyworm larva eats by remaining in the whorl of maize, avoiding contact with insecticides that have been administered. Therefore, several systemic insecticides were studied against the fall armyworm of maize.  At present, in countries such as India, which the pest recently invaded, since there are no recommended insecticides in place, the recommendation has been made by Central Insecticide Board and Registration Committee. The committee recommends the use of chlorantraniliprole 18.5 SC, thiamethoxam 12.6% + lambda-cyhalothrin 9.5% ZC, and spinetoram 11.7 SC for fall armyworm management. However, potential large-scale use of these chemical insecticides may cause hormesis effects, pest resurgence, and resistance development in target insect pests [85,86,87,88,89]. Additionally, these insecticides may have multiple negative impacts on human health and non-target organisms [90,91,92].

4. Biological Approaches

4.1. Biological Control

Biological control appears as a potential alternative to the chemical management of fall armyworm [93,94]. In various places, many natural enemies have been discovered to be related to this pest. Molina-Ochoa et al. [95] documented 150 species of parasitoids and parasites associated with fall armyworm from the Americas and the Caribbean basin. In India, Shylesha et al. [20] also reported egg, larval, and larval–pupal parasitoids and predators attacking different stages of this pest on maize. Among the predators, various ground beetles (Coleoptera: Carabidae); the striped earwig, Labidura riparia (Pallas) (Dermaptera: Labiduridae); the spined soldier bug, Podisus maculiventris (Say) (Hemiptera: Pentatomidae); and the insidious flower bug, Orius insidiosus (Say) (Hemiptera: Anthocoridae) were found to be effective against fall armyworm of maize as a means of biocontrol [96]. While predators have a significant impact on fall armyworm survival and development, parasitoids, which are more effective in causing mortality in fall armyworm populations, significantly outnumber predators. Solitary parasitoids of the Hymenoptera genera Chelonus and Campoletis were recovered from S. frugiperda larvae [97] and Trichogramma parasitoids could be high potential biocontrol agents for developing inundative biological control programs [98,99,100]. In Africa, five species of parasitoids were recorded from the fall armyworm in three East African countries in 2017 [84]. The Campoletis species is a larval endoparasitoid that plays an important role in regulating S. frugiperda [95]. The fall armyworm is a natural host of the parasitoid C. flavicincta, the larvae of the parasitoid. The larvae of S. frugiperda starts making its cocoon, which allows it to continue its development until its adult emergence. At this juncture, prime emphasis must be given to identifying the potential natural enemies and disease-causing pathogens for biological control of S. frugiperda. Biocontrol and biopesticides approaches are eco-friendly, sustainable, and appropriate alternatives to chemical insecticides [101]. These approaches form a strong base, and they are the key component of any integrated pest management program (IPM). Fall armyworm can effectively control insect populations by introducing naturally occurring disease-causing infections as natural regulatory agents [102]. Several microbial infections have been investigated in hopes of controlling fall armyworm numbers. More than 53 parasitic species, comprising 43 genera and 10 families, have been discovered as attacking fall armyworm [103]. Although no commercial Bt isolates have been created to combat fall armyworm, the Cry1F protein is thought to be more toxic to fall armyworm than any other Cry protein [104]. Likewise, many predators that attack fall armyworm eggs and larvae were also reported.
Insect pathogenic viruses are also one of the important biocontrol agents of FAW. There are different viruses known to infect FAW larvae, such as Ascoviruses, Baculoviruses, Densoviruses, Rhabdoviruses, and Partiti-like viruses [105]. Among these, Baculoviruses are more promising. The marketability of virus products for the management of various insect pests around the world has improved as a result of recent technological advancements. Spodoptera frugiperda multiple nucleopolyhedrovirus (SfMNPV), a baculovirus infecting S. frugiperda, has become commercially and registered in several countries for FAW management [106,107].
Understanding the ecological and toxicological connection between genetically modified cultivars (GMCs) and biological control agents is critical for GM cultivar compatibility and integrated pest resistance management strategies. To support this, Souza et al. [108] reported the efficiency of biological control for fall armyworm resistance to the protein Cry1F. They studied the search behaviour and predatory capabilities of Oriusinsidiosus (Say) (Hemiptera: Anthocoridae) and Doru luteipes (Scudder) (Dermaptera: Forficulidae) on fall armyworm eggs and caterpillars resistant to the expressed protein Cry1F [108].

4.2. Bird Perches

A diverse group of insectivorous birds has been found to feed on crop insect pests [109]. These insectivorous birds have been known to reduce the larval population by as much as 84% [110]. According to Jones et al. [111], many birds occurring in cropped fields actively forage for and consume caterpillars in crop vegetation. Black drongo, house sparrows, blue jays, cattle egret, rosy pastor, and mynah are frequent insectivorous birds that feed on a large number of lepidopteran insects among predatory birds. Insectivore birds show a strong tendency of attracting large size larvae [111]. In the case of environmentally friendly pest management, fall armyworm larvae that have escaped parasitism grow faster than those parasitized by Euplectrus wasps [111]. Boundary trees (such as fodder, fuelwood, and shelter trees) provide perches and roosts for birds and bats and also increase the structural variety of the farm ecosystem by providing shade and shelter [44]. These bird species are capable of extracting fall armyworm larvae from maize plant whorls and husks. Based on these results, it was known that insectivorous birds are also a potential tool for the ecofriendly and sustainable management of fall armyworm of maize. Under field conditions, insectivorous birds normally attract large larvae, although red-winged blackbirds were equally likely to eat parasitized and non-parasitized fall armyworm prey of the same body size [111]. Maize is a non-branching crop plant, which discourages birds from sitting and staying for extended periods. Therefore, suitable live bird perches in the maize ecosystem should be established. In order to facilitate bird visits, several fast-growing plants that provide rigid support for perching insectivorous birds from the vegetative stage until crop maturity should be grown in the maize field.

4.3. Use of Botanicals

In nature, many plants exhibit insecticidal properties. Preparations made out of those plant products are called botanical insecticides. These are insect poisons that are found in nature and are extracted or generated from plants or minerals. Increased interest in the eco-friendly management of pests in most edible crops led to exploring the use of botanicals in pest management. Botanicals employed for the management of pests are safe for humans and animals with reduced environmental impact. Fall armyworm tends to become highly destructive in maize cultivation; synthetic chemicals used in maize production greatly impair the quality of maize kernels and fodder values. Therefore, the utilization of botanicals is reported to be a potential strategy in pest management [112]. Similarly, many studies were conducted in various parts of the world reported that botanicals found to be an alternative and potential strategy in fall armyworm management [113]. The greatest strength of botanical extracts is their specificity, as most are essentially nontoxic and non-pathogenic to animals and humans [114]. Various plant species have shown insecticidal properties against fall armyworm; for example, extracts of neem, Azadirachta indica (Family: Meliaceae), Argemone ochroleuca Sweet (Family: Papaveraceae) [115], Boldo, Peumusboldus Molina (Family: Monimiaceae), Jabuticaba, and Myrciaria cauliflora (Mart.) O. Berg (Family: Myrtaceae).
There is sufficient evidence, as shown by the research findings, showing there are several prospects for the use of botanical extracts in the management of fall armyworm. However, due to several operational constraints, utilization of these potentials is limited [116,117].

This entry is adapted from the peer-reviewed paper 10.3390/agronomy12092150

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