BIOLOGICAL CONTROL OF Spodoptera frugiperda (J.E SMITH) (LEPIDOPTERA: NOCTUIDAE) GLOBALLY AND NEWLY NATURAL ENEMY ASSOCIATIONS IN ASIAN COUNTRIES: A REVIEW

INTRODUCTION

TheFall armyworm (FAW) or Spodoptera  frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), is an extremely destructive agricultural pest indigenous to the Americas (Goergen et al., 2016). It possesses the capacity to fly over many kilometres, rapid breeding, and attack 353 host species consisting of numerous plant families, about 76 (Adhikari et al., 2020). Therefore, it is regarded as one of the agile, highly polyphagous, and destructive noctuid moths in the agricultural fields (Nagoshi and Meagher, 2008).  Globally, this pest significantly reduces crop yields, Zea mays L. being the most susceptible plant in the Poaceae family (Rwomushana et al., 2018). Other than the Poaceae family, Amaranthaceae, Brassicaceae, Amaryllidaceae, Fabaceae, Malvaceae, and Solanaceae plant families are also vulnerable to S. frugiperda attack (Lu and Adang, 1996).

Farmers tend to apply insecticides considering the damage severity of the global crop production caused by insect attacks (Abang et al., 2021). Though it is a recommended practice under integrated pest management (IPM), it is not a long-term solution or nature-friendly remedy. Despite it is easy control method, its overuse might cause an undesirable negative impact on nature (Kumar et al., 2022).   Continuous application of insecticides over many years can cause the development of resistance and pose a substantial threat to beneficial insects and biodiversity (Yu et al., 2003). More than 29 insecticide active ingredients belonging to six modes of action groups have been ineffective due to the resistance developed by S. frugiperda (Wu et al., 2019), which include some extensively used insecticide categories such as carbamates, organophosphates, and pyrethroids (Yu, 1991). Moreover, S. frugiperda can develop resistance against transgenic maize plants expressing Cry1A.105 (MON89034), and the Cry1F gene expressing (TC1507) genetically modified maize varieties (Farias et al., 2014; Bernardi et al., 2017). Furthermore, in the Americas, S. frugiperda can also develop resistance against Bacillus thuringiensis (Bt) proteins (Hung, 2021). These challenges accelerate attempts to find sustainable and eco-friendly options to control S. frugiperda. Bio- control is one of the sustainable, eco-friendly, efficient, economical, and safe solutions in crop protection (Assefa and Ayalew, 2019). Therefore, identification of natural enemies is crucial to implement the bio-control management of S. frugiperda (Chhetri and Acharya, 2019; Lamsal et al., 2020).

Native natural enemies play a major roll to defending against the invasive pest species (Firake and Behere, 2020). Therefore, searching for native natural enemies against invasive S. frugiperda is crucial, and it is one of the initial steps of the pest management involving bio- control. Numerous bio-control agents of S. frugiperda have been found by Asian nations despite it recently invading Asia. Therefore, available literature on the bio-control agents of S. frugiperda in some countries of Asia is gathered in this review, along with the evidence of some successful attempts and their potential for the bio-control programme in some countries. For this purpose the scientific evidence from research papers published on the bio-control agents of S. frugiperda was carefully examined using various online databases, including Google Scholar, WoS, Scopus, PubMed, and ScienceDirect. All search terms included “fall armyworm,” or “FAW,” or “Spodoptera  frugiperda,” combined with following keywords: “natural enemies,” “Asia,” “predators,” “parasitoids,” “pathogens,” “entomopathogens,” “parasitism,” and “biological control.” Based on the relevance to the review paper, articles were selected. Ultimately, 130 of the most pertinent research papers were thoroughly reviewed to compile the content of the review paper. Additionally, the associations of natural enemies in Asian countries were also tabulated scientifically.

The dissemination of S. frugiperda in the Asian continent after America and Africa: S. frugiperda originated in America. Subsequently, in late 2016, it invaded the African continent (Cock et al., 2017). Afterward, it was dispersed to Asia in 2018 (Mahat et al., 2021). S. frugiperda was reported in South Asia in 2018, with reports from India (Sharanabasappa et al., 2018), subsequently reported in Sri Lanka (Perera et al., 2019), and Bangladesh (Alam et al., 2018). It was reported in Nepal (Guragain, 2019), Bhutan (Mahat et al., 2021), and Pakistan (Gilal et al., 2020) in 2019. Afterward, it invaded from South Asia to Southeast Asia rapidly (Lamsal et al., 2020). Consequently, it reported Myanmar and Thailand in 2018 as the first time in this region (IPPC, 2021). Subsequently, it was reported in Indonesia (Sartiami et al., 2020), Malaysia (Jamil et al., 2021), Vietnam (FAO, 2019d), the Philippines, and Laos (IPPC, 2021). It has been reported in East Timor, Cambodia, and Brunei by 2021 (IPPC, 2021). It was first reported in China as the first country in the East Asian region in 2018, and by 2019, it had spread to Japan and South Korea (IPPC, 2021).

Figure 1. The global dispersal of S. frugiperda. ((Source : FAO, 2020)

Opportunities for the bio-control of S. frugiperda: Bio-control refers to the beneficial impact of the predators, competitors, parasitoids, and pathogens in controlling pest populations (Nafiu et al., 2014). Natural enemies act as the bio-control agents in the biological control programme as a sustainable and cost-effective tool instead of synthetic chemicals (Ogunfunmilayo et al., 2021). Limited usage of pesticides by the farmers is a direct impact of the biological control, it enhances the food quality and food security, and ultimately it benefit to human health (Van Lenteren, 2012). Cock et al. (2010) specified that the biological control positively links with the ecosystem services, biodiversity conservation, and invasive species management.  However, to succeed in a bio-control pest management plan, identification of bio-control agents is vital (Bonsignore and Vacante, 2012a).The bio-control potential can be assessed from the global identification of their natural enemies. For instance, more than 150 parasitoid species associated with S. frugiperda have been reported in the Caribbean and the Americas (Molina-Ochoa et al., 2003a). The majority belongs to the orders Hymenoptera and Diptera, with Ichneumonidae and Braconidae being the widely distributed families within Hymenoptera, while Tachinidae was the widely distributed family in Diptera (Molina-Ochoa et al., 2003). About 30 native parasitoid species targeting S. frugiperda have already been discovered in 17 African countries (Sisay et al., 2019b). Additionally, numerous parasitoid species have been  identified predominantly in India and China, associated with S. frugiperda (Shylesha et al., 2018; Sharanabasappa et al., 2019; Yang et al., 2022).

Parasitoids exhibit a higher degree of specificity and are closely linked to one of the pest stages (Gowda et al., 2021). Egg parasitoids can control the pest before crop damage occurs. Therefore, they are recognized as the unique natural enemies for controlling agricultural pest species. During the parasitization process, they specifically search for the egg stage and attack at that stage. Consequently, they can break the further development and destroy the pest at the egg stage. Therefore, they are recognized as the efficient bio-control agents for the mass-reared and released strategy in a sustainable pest management approach (Parra and Coelho Jr. 2019). The genus Telenomus and Trichogramma are two of the prevalent egg parasitoid genera, including many egg parasitoid species that can parasitized the eggs of S. frugiperda successfully (Zang et al., 2021). Therefore, they are recognized as the ideal option for targeting S. frugiperda eggs in the biological pest management programme (Agboyi et al., 2020).  Telenomus remus (Nixon) is one of the efficient egg parasitoid species due to its high fecundity and its unique parasitism ability, because it can reach the bottom layers of the S. frugiperda egg mass and parasitize the whole egg mass (Bueno et al., 2008).  In contrast, Trichogramma species are partially effective because they cannot reach inside layers of S. frugiperda egg mass and are unable to parasitize the whole egg mass (Laminou et al., 2020). Other than egg parasitoids, an egg- larval parasitoid, Chelonus insularis (Cresson), is a frequently reported parasitoid species in the native range (Bahena and Cortez, 2015). Larval parasitoids, namely Campoletis sonorensis (Cameron), Campoletis flavicincta (Ashmead), and Pristomerus spinator (Fabricius) also abundantly recorded in the native range (Molina-Ochoa et al., 2003). In the African continent, the most prevalent larval parasitoids of S. frugiperda belong to the ichneumonid Charops sp., braconids Chelonus bifoveolatus (Szepligeti), C. curvi-maculatus (Cameron),  Cotesia icipe (Fernandez-Triana and Fiaboe), and tachinids Drino quadrinozula (Thomson) (Amadou et al., 2018; Agboyi et al., 2020;  Abang et al., 2021; Otim et al., 2021).

Not only parasitoids but also many predator species significantly contribute to managing S. frugiperda (Firake and Behere, 2020a). Some of them have been successfully employed under the augmentative release approach to efficiently control the pest populations (Collier and Van Steenwyk, 2004). Predators are capable of targeting multiple developmental stages of fall armyworm (Abbas et al., 2022), and different taxa specialize in attacking distinct life stages (Harrison et al., 2019).  Varella et al. (2015) who mentioned that the eggs and larvae of S. frugiperda are more vulnerable to predation by different predator species than to parasitoid attack. Bahena and Cortez (2015) documented sixty five predator species of S. frugiperda, primarily associated with the egg and larval stages. Several insect predators belonging to Dermaptera, Coleoptera, Hymenoptera, and Hemiptera have been effectively utilized in augmentative biological control programmes across the Americas (Abbas et al., 2022). Commonly reported predators include Orius tristicolor (White) and O. insidiosus (Say) (Hemiptera: Anthocoridae), Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), Sinea confusa (Caudell) (Hemiptera: Reduviidae), Doru lineare (Eschscholtz), and D. luteipes (Scudder) (Dermaptera: Forficulidae) (Bahena and Cortez, 2015; Varella et al., 2015). S. frugiperda larvae are also preyed on by certain social and solitary wasp species (Sousa et al., 2011; Southon et al., 2019). In India, twelve predator species were reported by Firake and Behere (2020a), including tiger beetles (Coleoptera: Cicindelidae), paper wasps (Hymenoptera: Vespidae), and earwigs (Dermaptera: Forficulidae), with the pentatomid bug Eocanthecona furcellata (Wolf) identified as the most dominant predator. Beyond insect predators, nematodes (Sun et al., 2020), various spider species (Anandhi and Saminathan, 2021), bacteria (Sivakumar et al., 2020), fungi (Manjula et al., 2019), and viruses (Raghunandan et al., 2019) also act as an integral part of the bio- control agents of S. frugiperda.

Successful evidence for bio-control implementation against S. frugiperda in the field: In the American continent, numerous bio-control agents, such as predators, pathogens,  and parasitoids of S. frugiperda have been documented (Molina-Ochoa et al., 2003). In African countries, they have also reported a wide range of bio-control agents after the invasion of S. frugiperda (Sisay et al., 2019). Though it has recently invaded Asia, some Asian countries reported many natural enemies of S. frugiperda within this time duration (Firake and Behere, 2020). According to Ahissou et al. (2021), S. frugiperda mortality on its native continent can reach 42% due to natural enemies, which significantly contributes to maintain the natural equilibrium of the pest population. Firake and Behere (2020a) reported that natural enemies in maize fields in northeastern India cause between 57 and 73% of S. frugiperda larval mortality. Unsprayed fields in America exhibit more than 44% of natural parasitism against S. frugiperda (FAO, 2017).

Telenomus remus was first introduced into the Americas through field releases in Barbados during 1971–1972, achieving parasitism rates of over 60% (Cave, 2000). Subsequent augmentative releases in Latin America and Venezuela have resulted in up to higher egg parasitism about 90% (Cave, 2000; Ferrer et al., 2001). Field evaluations further revealed parasitism levels of 80–100% in Venezuela and Ghana, providing effective suppression of S. frugiperda populations (Cave, 2000; Pomari et al., 2012). Native parasitoids from Sub-Saharan African countries exhibit higher levels of parasitism (Assefa and Ayalew, 2019). In Niger, experimental releases of T. remus alongside a Trichogrammatoidea sp. in sorghum fields achieved 64% parasitism in the initial field experiments (Laminou et al., 2020). Abang et al. (2021) reported that natural parasitism of egg parasitoids (T. remus and T. chilonis) ranged from 50 to 100% across various agro-ecological zones in Cameroon. Furthermore, T. remus demonstrated parasitism of eggs exceeding 50% in Tanzania and Kenya (Sisay et al., 2019). Sadore in Niger, the augmentative release of T. remus led to a 64% parasitism (Laminou et al., 2020), and between 60% and 90% in Latin America (Cave, 2000). According to Agboyi et al. 2021, field parasitism of T. remus was observed at 30 to 100% in Ghana.

In Karnataka, India, egg parasitism of S. frugiperda by Trichogramma chilonis (Ishii) was 24%, and by T. remus was 9% (Navik et al., 2021). When S. frugiperda first invaded in 2018, parasitism by T. chilonis ranged between 1.08 to 1.20% respectively.  However, in 2019-2020, parasitism level increased significantly, ranging from 2.28 to 20% across various regions in Karnataka, India. T. chilonis parasitism rates varied from 7.5 to 18% in Maharashta. T. remus parasitism in Karnataka was between 1.0% and 7% in 2018, and 1.2 to 8% in 2019-20 (ICAR-NBAIR, 2020). Hainan province in China, T. remus parasitized 28.9% of egg masses (Tang et al., 2020a), and in Guangdong province it was 30.6%   (Huo et al., 2019). In southeastern China, field application of T. remus to control FAW resulted in 100% parasitism for egg masses and 84% for individual eggs (Zhao et al., 2020). In maize fields of Shandong Province, China, studies on T. chilonis recorded 73% egg mass parasitism and 86% for individual eggs (Yang et al., 2019).

Other than egg parasitoids, larval parasitoids also contribute to considerably higher parasitism levels in the field. For instance, larval parasitism of S. frugiperda in the pesticide free fields in the southern USA has reached up to 44% (Meagher et al., 2016). In Costa Rica and Nicaragua, reported  more than 60% larval parasitism of S. frugiperda (Marenco and Sauders, 1993). In Ethiopia, Cotesia icipe (Hymenoptera: Braconidae) has shown 33.8 to 45.3% larval parasitism, being a predominant larval parasitoid in Ethiopia (Sisay et al., 2018).  In Kenya, the most prevalent Tachinid fly, Palexorista zonta exhibit parasitism rate of 12.5%, in Tanzania and Kenya, Charops ater and Coccygidium luteum parasitism rates are reported to be 4.0 to 8.3% and 6.0 to 12%, respectively (Sisay et al., 2018). Agboyi et al. (2021) stated that larval parasitoids severely attack young larvae of S. frugiperda in Ghana. Moreover, more than 20% of S. frugiperda larvae were parasitized by Coccygidium luteum (Brulle) in Ghana and Benin (Agboyi et al., 2020).Chelonus bifoveolatus (Szepligeti) was one of the predominant parasitoid species reported in West Africa (Koffi et al., 2020). Relatively low larval parasitism level reported in Uganda, averaging 9.2% (Otim et al., 2021). A similar result was reported in Mozambique, averaging 9.5% larval parasitism (Canico et al., 2020). Rate of parasitism of Che. bifoveolatus ranged from 0.8 to 16.7% in Uganda (Otim et al., 2021) it ranged from 0 to 35.6% in Ghana (Agboyi et al., 2020). In addition parasitism rate of Chelonus sp. was 10.9% in Senegal (Tendeng et al., 2019). Parasitism rate of Coc. luteum reported in Ethiopia, Tanzania, and Kenya were 4.6, 5.0, and 8.3%, respectively (Sisay et al., 2018).

The entomopathogens also play an imperative role to managing S. frugiperda. In Karnataka Metarhizium rileyi infections ranged from 10 to 62% across different states of natural field settings (Mallapur et al., 2018; Shylesha et al., 2018). ICAR-NBAIR (2020) recorded that Beauveria felina accounted for nearly 30% of natural S. frugiperda infections in Chikkaballapur, Karnataka.Several isolates of S.  frugiperda nucleopolyhedrovirus (SfNPV) have been employed worldwide for the bio-control of S. frugiperda, achieving control efficacies above 80%(Gomez et al., 2013; Behle and Popham, 2012).

Bio-control agents of S. frugiperda in some Asian countries: Diverse groups of bio-control agents have been documented  in Asian nations, including India (Shylesha et al., 2018), China (Tang et al., 2021), the Philippines (Navasero and Navasero, 2020), Pakistan(Riaz et al., 2024), Indonesia (Tawakkal et al., 2021), Malaysia (Singhamuni et al., 2025), Sri Lanka (Bandara et al., 2021) and Nepal (Elibariki et al., 2020) such as parasitoids, predatory insects, spiders, nematodes, and entomopathogens. Following S. frugiperda's invasion as an agricultural pest in Asia, many scientists have studied and reported the diversity of its natural enemies, which are tabulated herein.

Parasitoids: Parasitoids represent a diverse group of natural enemies. In Asian countries, more than 50 parasitoid species have been documented in association with S. frugiperda (Table 1).

Table1. Parasitoids associated with S. frugiperda in Asian countries

Predators: Predators are capable of attacking various developmental stages of S. frugiperda (Abbas et al., 2022). Numerous predators including insects and spiders have been documented throughout Asia (Table 2).

Table 2. Predatory insects and spider species associated with S. frugiperda in Asian countries

Nematodes: Nematodes play an ecologically beneficial role and are essential for managing soil-dwelling insect pests, particularly pupae of the armyworms (Dillman et al., 2012). S. frugiperda pupae are highly susceptible to these entomopathogenic nematodes, with a reported infestation potential of approximately 23,000 per square foot (Prasanna et al., 2018).

Table3. Parasitic nematodes associated with S. frugiperda in Asian countries

Entomopathogens (Fungi, Bacteria, Viruses): Insect diseases are caused by entomopathogens, primarily bacteria, fungi, and viruses. Entomopathogenic fungi generally target S. frugiperda larvae and pupae. Through an insect's integument, fungal spores penetrate and develop within the body. After multiplying, they release specific toxins that degrade tissues and kill the insect (Abbas et al., 2022). According to Sujeetha and Sahayaraj (2014), when an insect is infected by entomopathogenic fungi, it ceases feeding, exhibits colour changes such as brown, reddish, green, or cream depending on the fungal species involved, and eventually dies. In natural fields, the pest acquires the virus by consuming contaminated maize leaves (CABI, 2021). These virions start to replicate in the nucleus after infecting the midgut's epithelial cells (Prasanna et al., 2018). Additionally, after spreading throughout the body cavity, these viruses begin to infect other tissues. The skin discoloration, blemishes, and decreased feeding are the primary symptoms of a Baculovirus infection (Valicente, 1988). The infected larvae are dark-colored, soft, and have secretions rich in polyhedrons, promoting the virus's propagation (Valicente, 1988).

Five entomopathogenic fungi species, one bacterial species, and two viruses were recorded as entomopathogens of S. frugiperda larvae and pupae in Asian countries (Table 4), with entomopathogenic fungi being the most widely distributed. Metarhizium and Beauveria species are the dominant types. Bacillus thuringiensis ranks as one of the most frequently used entomopathogenic bacteria against insects globally. It is also found in India, attacking S. frugiperda (Firake and Behere, 2020a; Sivakumar et al., 2020). Sf NPV represents one of the dominant viruses attacking S. frugiperda in Asia (Raghunandan et al., 2019).

Table 4. Entomopathogens associated with S. frugiperda in Asian countries

Significance of the conservation of natural enemies: These findings highlight the presence and vital role of indigenous bio-control agents in suppressing invasive species, acting as the first line of defence. This occurs because locally available bio-control agents of closely linked pest species often broaden the host range and contribute substantially to the control of invasive pests naturally (Vercher et al., 2005). The existence of native natural enemies associated with similar pest species, therefore, represents the initial protective barrier against new invasions. Hence, empowerment of the farmers with positive attitudes regarding native natural enemies to control invasive pest species is crucial for biodiversity conservation in agricultural fields. Evaluating the natural control inflicted by native bio-control agents is a significant approach in implementing a combined management approach for invasive insect pests (Firake and Behere, 2020a). Consequently, studies on local bio-control agents in newly invaded regions are of great importance.

Many biological control treatments are unacceptable to farmers because they require immediate solutions to pest problems in their cultivated fields (Ahissou et al., 2021). Therefore, most farmers tend to find quick methods other than bio-control. They do not much consider the sustainable management of pests in their field. A key limitation of biological control is that it acts more slowly than synthetic pesticides (Rioba and Stevenson, 2020). Nevertheless, biological control plays an essential role in conserving biodiversity while offering significant economic benefits (Epstein et al., 2021). As with traditional pest control methods, evaluating the effectiveness of bio-control agents must account for long-lasting effects in addition to short-terms outcomes (Ahissou et al., 2021).

Natural enemies provide an effective means of controlling S. frugiperda. Yet, many farmers remain unaware of their existence, and indiscriminate insecticide use often diminishes their populations. Since most farmers rely primarily on chemical control, enhancing their knowledge and attitudes toward natural enemies through targeted extension and education programmes is essential.

Conclusion: According to the literature survey, locally available natural enemies in various Asian countries have successfully established new associations with S. frugiperda. The literature review indicates that about 55 species of parasitoids, 28 species of predatory insects, 6 species of spiders, 4 species of nematodes, 7 species of entomopathogenic fungi, two species of entomopathogenic bacteria, and two species of entomopathogenic viruses have been recorded in China, India, Pakistan, Indonesia, Sri Lanka, the Philippines, Nepal, and Malaysia a few years after the invasion of S. frugiperda. These findings highlight the critical role that local natural enemies play in combating invading pest species, acting as the forefront of defence. This is because the locally available bio-control agents targeting closely linked pest species often expand their feeding range and contribute significantly to the natural management of invading pests. This review highlights the natural enemies that have established novel associations with S. frugiperda in several Asian countries, evaluates their potential as bio-control agents, and summarizes evidence of their successful application in globally. We hope that this synthesis of knowledge will encourage other affected Asian nations to adopt ecologically friendly suppression of S. frugiperda.

Acknowledgement:  The authors are indebted to the Kementerian Pengajian Tinggi for awarding the scholarship (MIS-KPT.B(S) 700-4/2/1JLD.4 (84).

REFERENCES

Abang, A. F., S. N. Nanga, A. Fotso Kuate, C. Kouebou, C. Suh, C. Masso and K. K. M. Fiaboe (2021). Natural enemies of fall armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) in different agro-ecologies. J. Insects. 12(6): 509. https://doi.org/10.3390/insects12060509.

Abbas, A., F. Ullah, M. Hafeez, X. Han, M. Dara, H. Gul and C.R Zhao. (2022). Biological Control of fall armyworm, Spodoptera frugiperda. J. Agron. 12 (11): 2704.https://doi.org/10.3390/agronomy12112704.

Adhikari, K., S. Bhandari, L. Dhakal and J. Shrestha (2020). Fall armyworm (Spodoptera frugiperda) A threat in crop production in Africa and Asia. Peruv. J.Agron. 4(3): 121-133.https://doi.org/10.21704/pja.v4i3.1495.

Agboyi, L. K., B. F. R. Layode, K.O. D. Fening, Babendries et al. (2021). Assessing the potential of field releases of Telenomus remus to control Spodoptera frugiperda in Ghana. J. Insects 12(8): 665. https://doi.org/10.3390/insects 12080665.

Agboyi, L. K., G. Georgen, P. Beseh, S. A. Mensah and M. Kenis (2020). Parasitoid complex of fall armyworm, Spodoptera frugiperda in Ghana and Benin. J. Insects. 11(2):68. https://doi.org/ 10.3390/insects11020068.

Ahissou, B. R., W.M. Sawadogo, A.H. Bokonon-Ganta, I. Somda and F. J. Verheggen (2021b). Integrated pest management options for the fall armyworm Spodoptera frugiperda in West Africa: Challenges and opportunities. A review. Biotechnol. Agron. Soc. Environ. 25(3): 192–207. https://doi.org/10.25518/1780-4507.19125.

Alam, S. N., D. Sarker, M.Z.H. Pradhan, M.H. Rashid, M.A. Sarkar, K. Begum and M.A Mannan (2018). First report of occurrence of fall armyworm Spodoptera frugiperda in Bangladesh. Bangladesh. J. Entomol. 28: 97–101.

Amadou, L., I. Baoua, M. N. Ba, L. Karimoune and R. Muniappan (2018). Native parasitoids recruited by the invaded fall army­worm in Niger. Indian J. Entomol. 80(4): 1253–1254. https://doi.org/10.5958/0974-8172.2018.00338.3.

Anandhi, S. and V. R. Saminathan (2021). New record of larval parasitoids and predatory spiders on fall armyworm Spodoptera frugiperda (JE Smith) (Noctuidae: Lepidoptera) in Tamil Nadu. J. Entomol. Zool. Stud. 9(4): 340-342.

Assefa, F. and D. Ayalew (2019). Status and control measures of fall armyworm (Spodoptera frugiperda) infestations in maize fields in Ethiopia. Cogent Food Agric. 5(1): 1641902. https://doi.org/10.1080/23311932.2019.1641902.

Bahena, J. F. and M. E. Cortez (2015). Fall armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae). Biological control cases in Mexico. Basic Agricultural library; Guadalajara (Mexico). 181–250 pp.

Bandara, K. A. N. P., S. S. Weligamage, K. M. D. W. P. Nishantha and Y. D. G. C. Yubak (2021). Sustainable Management of Fall Armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae) in Maize Cultivation of Sri Lanka Through Local Natural Enemies. Trop. Agriculturist. 169(4): 33-41.https://doi.org/10.4038/ta.v169i4.5.

Behle, R.W. and H.J. Popham (2012). Laboratory and field evaluations of the efficacy of a fast-killing baculovirus isolate from Spodoptera frugiperda. J. Invertebr. Pathol. 109: 194-200. https://doi.org/10.1016/j.jip.2011.11.002.

Bernardi, D., O. Bernardi, R. Horikoshi, E. Salmeron, D. Okuma, J. Farias,  et al. (2017). Selection and characterization of Spodoptera frugiperda (Lepidoptera: Noctuidae) resistance to MON 89034xTC1507 xNK603 maize technology. Crop. Prot, 94 :64-68.https://doi.org/10.1016/j.cropro.2016.11.026.

Bonsignore, C.P. and V.Vacante (2012a). Natural enemies, Integrated Control of Citrus Pests in the Mediterranean Region. Bentham Science Publishers; London, pp. 66–87 eISBN: 978-1-60805-294-3.

Bueno, R.C.O.F., A.F. Bueno, T. R. Carneiro, D. Pratissoli and O.A. Fernandes (2008). Biology and thermal requirements of Telenomus remus Nixon (Hymenoptera: Scelionidae) reared on fall armyworm Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) eggs. Cienc Rural. 38:1–6. https:// doi. org/ 10. 1590/ S0103-​84782 00800 01000 01.

CABI (2021). Spodoptera frugiperda. In Invasive Species Compendium. UK; Wallingford. CAB International. https://doi.org/10.1079/ISC.29810.20203373913.

Caniço, A., A. Mexia and L. Santos (2020). First report of native parasitoids of fall armyworm Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) in Mozambique. J. Insects. 11(615): 1-12. https:// doi.org/10.3390/insects11090615.

Cave, R. D. (2000). Biology, ecology and use in pest management of Telenomus remus. Biocontrol News Inf. 21: 21–26.

Chhetri, L. and B. Acharya (2019). Fall armyworm(Spodoptera frugiperda): A threat to food security for south Asian Country. Control and management options. FAM. 4 (1): 38-44.https://doi.org/10.31830/2456-8724.2019.004.

Cock, M., P. Beseh, A. Buddie, G. Cafa and J. Crozier (2017). Molecular methods to detect Spodoptera frugiperda in Ghana and implications for monitoring the spread of invasive species in developing countries. Sci. Rep. 7(1): 4103.https://doi.org/10.1038/s41598-017-04238-y.

Cock, M.J.W., J.C. van Lenteren, J. Brodeur, B.I.P. Barratt, F. Bigler, K. Bolckmans, F.L. Consoli, F. Haas, P.G. Mason and J.R.P. Parra (2010). Do new access and benefit sharing procedures under the convention on biological diversity threaten the future of biological control? BioControl. 55(2): 199–218. https://doi.org/ 10.1007/s10526-009-9234-9.

Collier, T. and R. van Steenwyk (2004). A critical evaluation of augmentative biological control. Biol. Control. 31: 245–256. https://doi.org/10.1016/j.biocontrol.2004.05.004

Dillman, A. R., J.M. Chaston, B.J. Adams, T.A. Ciche, H. Goodrich-Blair, S.P. Stock and P.W. Sternberg (2012). An entomopathogenic nematode by any other name. PLoS Pathogens, 8(3): e1002527. https://doi.org/10.1371/journal.ppat.1002527.

Elibariki, N., A.S.R. Bajracharya, B. Bhat, T. Tefera, J.L.  Mottern, G. Evans and P. Likhay (2020). Candidates for augmenta­tive biological control of Spodoptera frugiperda (JE smith) in Kenya, Tanzania and Nepal. Indian J. Entomol. 82(4): 606–608. https://doi.org/10.5958/0974-8172.2020.00088.7.

Epstein Y., G. Chapron and F. Verheggen (2021). EU court to rule on banned pesticide use. J. Science, 373(6552): 290.  https://doi.org/10.1126/science.abj9226.

FAO (2017). Sustainable management of the fall armyworm (Spodoptera frugiperda) in Africa: A framework for partnership. Rome, Italy: Food and Agriculture Organization of the United Nations. Retrieved December, 2024, from 
https://www.fao.org/fileadmin/templates/fcc/map/map_of_affected_areas/Fall_Armyworm_brief_-__24Nov17_.pdf.

FAO (2019d). Fall armyworm reported in Vietnam. Food and Agriculture Organization of the United Nations (FAO) Retrieved October, 2024, from.
https://www.fao.org/fall-armyworm/background/overview/en/.

FAO. (2020). The Global Action for Fall Armyworm Control: Action framework 2020-2022. Working together to tame the global threat. Rome: Food and Agriculture Organization of United Nation. Retrieved December, 2024, from https://openknowledge.fao.org/server/api/core/bitstreams/e50d4cd9-c774-4a3b-ab8e-6144eca8af5c/content.

Farias, J., D. Andow, R. Horikoshi, R. Sorgatto, P. Fresia, A Dos Santos, et al. (2014). Field- evolved resistance to Cry1F maize by Spodoptera frugiperda (Lepidoptera:Noctuidae) in Brazil. Crop. Prot, 64: 150-158.https://doi.org/10.1016/j.cropro.2014.06.019.

Ferrer, F., S. Biologico, C. Antigua Yaritagua-Barquisimeto, S. Chorobobo and E. Lara (2001). Biological control of agricultural insect pests in Venezuela; advances, achievements, and future perspectives. Biocontrol News Inf.  22: 67–74.

Firake, D. M. and G.T. Behere (2020a). Natural mortality of invasive fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae) in maize agroecosystems of north­east India. Biol. Control. 148: 104303. https://doi.org/10.1016/j.biocontrol.2020.104303.

Firake, D. M. and G.T. Behere (2020b). Bioecological attributes and physiological indices of invasive fall armyworm, Spodoptera frugiperda (J. E. Smith) infesting ginger (Zingiber officinale Roscoe) plants in India. J. Crop Prot. 137: 105233. https://doi.org/10.1016/j.cropro.2020.105233.

Ghosh, E., R. Varshney and R. Venkatesan (2020). Performance of Bracon brevicornis (Wesmael) on two Spodoptera species and application as potential biocontrol agent against fall armyworm. Bio.Rxiv. 2020:06.https://doi.org/10.1101/2020.06.27.171025.

Gilal, A. A.,  L. Bashir, M. Faheem, A. Rajput, J.A. Soomro, S. Kunbhar  and  J.G.M. Sahito  (2020). First record of invasive fall armyworm (Spodoptera frugiperda (Smith) (Lepidoptera: Noctuidae)) in corn fields of Sindh, Pakistan. Pak. J. Agric.Res. 33(2): 247–252. http://dx.doi.org/10.17582/journal.pjar/2020/33.2.247.252

Goergen, G., P. Kumar, S. Sankung, A. Togola  and M. Tam ( 2016). First report of outbreaks of the fall armyworm Spodoptera frugiperda (J.E.Smith)(Lepidoptera,Noctuidae),a new alien invasive pest in West and Central Africa. PLoS One 11 (10): e0165632. https://doi.org/10.1371/journal.pone.0165632.

Gomez, J., J. Guevara, P. Cuartas, C. Espinel and L. Villamizar (2013). Microencapsulated Spodoptera frugiperda nucleopolyhedro virus: insecticidal activity and effect on arthropod populations in maize. Biocontrol Sci. Technol. 23: 829-846. https://doi.org/10.1080/09583157.2013.802288.

Gupta, A. and A. N. Shylesha (2021). Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae), another addition to the native parasitoid complex of the fall armyworm, Spodoptera fru­giperda (J. E. Smith) (Lepidoptera: Noctuidae) in India. J. Biol. Control. 35(2): 127–129. https://doi.org/10.18311/jbc/2021/28119.

Gupta, A., P.L. Soujanya, C. van Achterberg and J.C. Sekhar (2020b). Coccygidium transcaspicum (Kokujev) (Hymenoptera: Braconidae) parasitizing larvae of invasive pest Spodoptera fru­giperda (J. E. Smith) (Lepidoptera: Noctuidae) in India. Zootaxa, 4750(2): 293–297. https://doi.org/10.11646/zootaxa.4750.2.13.

Gupta, A., Y. Lalitha, R. Varshney, A.N. Shylesha and C. Van Achterberg (2020a). Chelonus formosanus Sonan (Hymenoptera: Braconidae) an egg-larval parasitoid of the invasive pest Spodoptera frugiperda (JE Smith)(Lepidoptera: Noctuidae) amenable to laboratory mass production in India. J. Entomol. Zool. Stud. 8(1): 1521-1524.

Guragain, M. (2019). Lab report confirms entry of American fall armyworm in Nepal. Retrieved June, 2024, from, https://myrepublica. nagariknetwork. com/news/lab-reportconfirmsentry-of-american-fall-armyworm-in-nepal.

Harrison, R. D., C. Thierfelder, F. Baudron, P. Chinwada, C. Midega, U. Schaffner and J. van den Berg (2019). Agro-ecological options for fall armyworm (Spodoptera frugiperda JE Smith) management: Providing low-cost, smallholder-friendly solutions to an invasive pest. J. Environ. Manage. 243: 318–330. https://doi.org/10.1016/j.jenvman.2019.05.011.

Hou, Z. R., B.B. Sun, X.J. Liu, Z. Yin, J.  Li and X. Guo (2020). Predatory functional response of assassin bug Sycanus falleni to the larvae of fall armyworm Spodoptera frugiperda. J.  Plant Prot. 47(4): 852-858. https://doi.org/ 10.13802/j.cnki.zwbhxb.2020.2020823.

Hui, L. I., S.S. Jiang, H.W. Zhang, G.E.N.G. Ting, K.A. Wyckhuys and K.M. WU (2021). Two-way predation between immature stages of the hoverfly Eupeodes corollae and the invasive fall armyworm (Spodoptera frugiperda JE Smith). J. Integr. Agric. 20(3): 829-839. https://doi.org/ 10.1016/S2095‑3119(20)63291‑9.

Hung, F (2021). Resistance of the fall armyworm, Spodoptera frugiperda to transgenic Bacillus thuringiensis Cry 1F corn in the Americas: lessons and implications for Bt corn IRM in China. Insect Sci. 28: 574-589.https://doi.org/10.1111/1744-7917.12826.

Huo, L. X., J.C. Zhou, S.F. Ning, Q. Zhao, L.X. Zhang, Z.T. Zhang, L.S. Zhang and H. Dong (2019). Biological character­istics of Telenomus remus against Spodoptera frugiperda and Spodoptera litura eggs. In Chinese. Plant Prot. 45: 60–64. https://doi.org/ 10.16688/j.zwbh.2019406.

ICAR-NBAIR (2020). Annual Report 2019. ICAR-National Bureau of Agricultural Insect Resources, India; Bengaluru, 105 pp.

IPPC (2021). Official pest reports. Retrieved June, 2024, from, https://www.ippc.int/en/coun­tries/all/pestreport/.

Jamil, S., M. Saranum, L. Hudin and W. Ali (2021). First incidence of the invasive fall armyworm Spodoptera frugiperda (smith.J.E,1797) attacking maize in Malaysia. Biolnvasions Rec. 10 (1): 81-90. https://doi.org/10.3391/bir.2021.10.1.10.

Jindal, J., K.P. Sharma, P.S. Shera and H.K. Cheema (2022). Native Parasitoids of Fall Army Worm Spodoptera frugiperda (JE Smith) in Maize. Indian J.  Entomol. 84(4): 865-867.

Jin, T., Y. Lin, G. Ma, J. Liu, Z. Hao, S. Han and Z. Peng (2021). Biocontrol potential of Trichogramma species against Spodoptera frugiperda and their field efficacy in maize. Crop Prot. 150, 105790.https://doi.org/10.1016/j.cropro.2021.105790.

Keerthi, M. C., A. Sravika, H. S. Mahesha, A. Gupta, H.A. Bhargavi and S. Ahmed (2020). Performance of the native predatory bug, Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae), on the Fall Armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera:Noctuidae), and its limitation under field condition. Egypt J. Biol. Pest Control. 30: 1-4. https://doi.org/ 10.1186/s41938‑020‑00272‑7.

Keerthi, M. C., S. S. Suroshe, S. Doddachowdappa, K.T. Shivakumara, H.S. Mahesha, V.S. Rana, A. Gupta, A. Murukesan, R. Casini, H.O. Elansari and N.A. Shakil (2023). Bio-Intensive Tactics for the Management of Invasive Fall Armyworm for Organic Maize Production. J. Plants. 12(3): 685.https://doi.org/10.3390/plants12030685.

Koffi, D., R. Kyerematen, V.Y. Eziah, K. Agboka, M. Adom, G. Goergen and R.L. Meagher (2020). Natural Enemies of the Fall Armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) in Ghana. Fla. Entomol. 103(1): 85–90. https://doi.org/10.1653/024.103.0414.

Kumar, R. M., B.G. Gadratagi, V. Paramesh, P. Kumar, Y. Madivalar, N. Narayanappa and F. Ullah  (2022). Sustainable management of invasive fall armyworm, Spodoptera frugiperda. J. Agron. 12(9): 2150.https://doi.org/10.3390/agronomy12092150.

Laminou, S. A., M. N. Ba, L. Karimoune, A. Doumma and R. Muniappan  (2020). Parasitism of locally recruited egg parasitoids of the fall armyworm in Africa. J. Insects. 11(7): 430. https://doi.org/10.3390/insects11070430.

Lamsal, S., S. Sibi  and S. Yadav (2020). Fall Armworm in South Asia: Threats and management. Asian J. Adv. Agric. Res. 21-34.https://doi.org/10.9734/AJAAR/2020/v13i330106.

Li, F., L. Wang, B. Lu, F. Cao, X. Pan, L. Yuan and S. Wu (2019). The report of Chelonus munakatae parasitizing fall armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) in Hainan, China. Chin. J. Biol. Control.  35 (6): 992–996.https://doi.org/10.16409/j.cnki.2095-039x.2019.06.003.

Li, W., Y. Jiang, M. Zhou, J. Chen and J. Yuan (2024). Stability of Spodoptera frugiperda multiple nucleopolyhedrovirus Hub1 and its field efficacy against S. frugiperda. Chin. J. Pest. Sci.  26(5), 956-961. https://doi.org/ 10.16801/j.issn.1008‑7303.2024.0068.

Liao, Y. L., B.Yang, M.F. Xu, W. Lin, D.S. Wang, K.W. Chen and H.Y. Chen (2019). First report of Telenomus remus parasitizing Spodoptera frugiperda and its field parasitism in southern China. J. Hymenop. Res. 73: 95-102. https://doi.org/ 10.3897/jhr.73.39136.

Lu, Y. and M. Adang (1996). Distinguishing fall armywarm (Lepidoptera:Noctuidae) strains using a diagnostic mitochondrial DNA marker. Fla. Entomol. 79: 48-55. https://doi.org/10.2307/3495753.

Mahat, K., A. Mitchell and T.  Zangpo (2021). An updated global COI barcode reference data set for fall armyworm (Spodoptera frugiperda) and first record of this species in Bhutan. J. Asia Pac. Entomol. 24(1): 105–109. https://doi.org/10.1016/j.aspen.2020.11.013.

Mallapur, C.P., A.K. Naik, S. Hagari, T. Praveen, P.K. Patil and S. Lingappa (2018). Potentiality of Nomuraea rileyi (Farlow) Samson against the fall armyworm, Spodoptera frugiperda (J E Smith) infesting maize. J. Entomol. Zool. Stud. 6(6): 1062-1067.

Manjula, K., Y.P. Saheb, M.J. Sudheer and A.R. Rao (2019). Studies on biology, feeding habits and natural enemies of fall armyworm, Spodoptera frugiperda, a new invasive pest in India. J. Entomol. Zool. Stud. 7(6): 1245-1250.

Marenco, R. A. and J.L. Saunders (1993). Parasitoides of Fall armyworm, Spodoptera frugiperda (Lepdidoptera: Noctuidae) in corn Turrialba Costa Rica. Integr. Pest Manag. 8 :13–18.

Meagher, R. L., G. S. Jr. Nuessly, R. N. Nagoshi and M. M. Hay-Roe (2016). Parasitoids attacking Fall Armyworm (Lepidoptera: Noctuidae) in sweet corn habitats. Biocontrol. 95: 66–72. https://doi.org/10.1016/j.biocontrol.2016.01.006.

Molina-Ochoa, J., J. E.  Carpenter, E. A. Heinrichs and J.E. Foster (2003a). Parasitoids and parasites of Spodoptera frugiperda (Lepidoptera: Noctuidae) in the Americas and Caribbean basin: an inventory. Fla. Entomol. 86(3): 254–289. https://doi.org/10.1653/0015-4040(2003)086[0254:PAPOSF]2.0.CO;2.

Molina-Ochoa, J., R. Lezama-Gutierrez, M. Gonzalez-Ramirez, M. Lopez-Edwards, M. A Rodriguez-Vega and F. Arceo-Palacios (2003b). Pathogens and parasitic nematodes associated with populations of fall armyworm (Lepidoptera: Noctuidae) larvae in Mexico. Fla. Entomol. 86(3): 244–253. https://doi.org/10.1653/0015-4040(2003)086[0244:PAPNAW]2.0.CO;2.

Nafiu, B.S., H. Dong and S. Mustapha (2014). Biological control of insect pests in West Africa: a review. Int. J. Appl. Res. Technol. 3(9): 39-45. https:// doi.org/10.13140/ RG.2.2.13034.31687.

Nagoshi, R. and R. Meagher (2008). Review of fall armyworm (Lepidoptera: Noctuidae) genetic complexicy and migration. Fla. Entomol. 91(4): 546-554.https://doi.org/10.1653/0015-4040-91.4.546.

Navasero, M. M. and M.V. Navasero (2020). Life cycle, morphometry and natural enemies of fall armyworm, Spodoptera frugiperda (JE Smith)(Lepidoptera: Noctuidae) on Zea mays L. in the Philippines. J. Int. Soc. Southeast Asian Agric. Sci. 26(2): 17-29.

Navik, O., A.N. Shylesha, J. Patil, T. Venkatesan, Y. Lalitha and T.R. Ashika (2021). Damage, distribution and natural enemies of invasive fall armyworm Spodoptera frugiperda (J. E. Smith) under rainfed maize in Karnataka, India. Crop. Prot. 143 :105536. https://doi.org/10.1016/j.cropro.2021.105536.

Ning, S. F., J.C.  Zhou, Z.T. Zhang, Q.J. Dong, X.P. Bai, L.S. Zhang  and H. Dong (2019). Five parasitic natural enemies of Spodoptera frugiperda and two hyperparasitoids of Cotesia glomerata were found in southeast of Guizhou province. J. Plant Prot. 45(6): 39-42. https://doi.org/ 10.16688/j.zwbh.2019454.

Ogunfunmilayo, A. O., S. A.  Kazeem, O.B. Adediby, L.C. Offord and T.I. Ofuya (2021). Occurrence of natural enemies of Spodoptera frugiperda in Nigeria. PLoS ONE, 16(7): e0254328. https://doi.org/10.1371/journal.pone.0254328.

Otim, M. H., S. A. Aropet, M. Opio, H.N. Opolot and W.T. Tay (2021). Parasitoids distribution and parasitism of Spodoptera frugiperda in different maize producing regions of Uganda.J. Insects. 12:121. https;//doi.org/10.3390/insects12020121.

Parra, J.R.P. and A. Jr. Coelho (2019). Applied biological control in Brazil: from laboratory assays to field application. J Insect Sci. 19:1–6. https:// doi. org/ 10.1093/ jisesa/ iey112.

Pebriansyah ( 2023). Predation power of Sycanus dichotomus Dohrn (Hemiptera : Reduvidae) on grayak caterpillar pest larvae Spodoptera frugiperda J.E Smith (Lepidoptera : Noctuidae) in the laboratory. Retrieved August, 2024, from https://repository.uisu.ac.id/bitstream/123456789/2553/1/Cover%2CBibliography.pdf.

Perera, N., M. Magamage, A. Kumara, H. Galahitigama, K. Dissanayake, C. Wekumbura, P. Iddamalgoda,C. Siriwardhana and P. Yapa (2019). Fall army­worm (FAW) epidemic in Sri Lanka: Ratnapura district perspec­tives. Int. J. Entomol. Res. 7(1): 9–18. https://doi.org/10.33687/entomol.007.01.2887.

Pomari, A. F., A. D. F. Bueno, R. C. O. De Freitas Bueno  and A. De Oliveira Menezes Junior (2012). Biological characteristics and thermal requirements of the biological control agent Telenomus remus (Hymenoptera: Platygastridae) reared on eggs of different species of the genus Spodoptera (Lepidoptera: Noctuidae). Ann.  Entomol. Soc. Am. 105(1): 73-81. https://doi.org/10.1603/AN11115.

Prasanna, B., J. Huesing, R. Eddy and V. Peschke (2018). Fall Armyworm in Africa: A Guide for Integrated Pest Management; USAID; CIMMYT, Mexico; Mexico City.

Qin, J., W. Qin, H. Chen, X. Cai, Y. Yu and H. Chen (2021). Two larval parasitoid wasps species of Spodoptera frugiperda (lepi­doptera: Noctuidae) found in Guangxi, China. J.Plant Prot. 47: 292–296.

Raghunandan, B. L., N. M. Patel, H.J. Dave and D.M. Mehta (2019). Natural occurrence of nucleopolyhedrovirus infecting fall armyworm, Spodoptera frugiperda (JE Smith)(Lepidoptera: Noctuidae) in Gujarat, India. J. Entomol. Zool. Stud.  7(2): 1040-1043.

Riaz, S., M. Ishtiaq, F.Z.A. Khan, G. Ali, M.A. Mehmood and M.S.Q. Zaman (2024). Occurrence of natural enemies in maize and the predatory potential of selected arthropods against fall armyworm in Multan, Pakistan. Int. J. of Trop. Insect Sci. 44(3): 1297-1307. https://doi.org/10.1007/s42690-024-01227-3.

Rioba, N. B and P.C. Stevenson (2020). Opportunities and scope for botanical extracts and products for the management of fall armyworm (Spodoptera frugiperda) for smallholders in Africa. Plants. 9(2): 207. https://doi.org/10.3390/plants9020207.

Rwomushana, I., M. Bateman, T. Beale, P. Beseh, K. Cameron, M. Chiluba et al. (2018). Research gate. from Fall armyworm: impacts and implications for Africa:Evidence Note update, Access date: 02-04-2023. https://www.invasive-species.org/wp-co ntent/uploads/sites/2/2019/02/FAW-Evidence-Note-October-2018.pdf.

Sagar, D., S.S. Suroshe, M. C.  Keerthi and R. Kumar (2022). Native parasitoid complex of the invasive Spodoptera frugiperda from Northern India. Int. J. Trop. Insect.Sci. 42: 2773-2778. https://doi.org/10.1007/s42690-022-00743-4.

Sari, L., N. Maryana and P. Hidayat (2021). Biology and life table of Trichogramma chilotraeae, Egg parasitoids of Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae). In IOP Conference Series: Environ. Earth Sci.  948 (1): 012044. IOP Publishing. https://doi.org/ 10.1088/1755-1315/948/1/012044.

Sari, W., N. Nelly, Hidrayani and Yaherwandi (2023). Natural enemies of Spodoptera frugiperda JE Smith (Lepidoptera: Noctuidae) on corn plants in West Sumatera. In IOP Conference Series: Environ. Earth Sci. 1160 (1): 012045. IOP Publishing.https://doi.org/ 10.1088/1755-1315/1160/1/012045.

Sari. A., D. Buchori and I. Nurkomar (2020). The potential of Telenomus remus Nixon (Hymenoptera: Scelinoidae) as biocontrol agent for the new fall armyworm S. frugiperda (Lepidoptera: Noctuidae) in Indonesia. Planta Tropika. 8(2): 69-74.https://doi.org/10.18196/pt.2020.116.-69-74.

Sartiami, D., I. Dadang Harahap, Y. Kusumah  and R. Anwar  (2020). First record of fall armyworm (Spodoptera frugiperda) in Indonesia and its occurence in three provinces. IOP Conf.Ser.Earth Environ. Sc, 468: 012021.https://doi.org/10.1088/1755-1315/468/1/012021.

Sharanabasappa, D. S., S. Kiran, A.  Naskar, P. Pradeep, C.M. Kalleshwaraswamy and K.N. Sharath (2021). First record of a parasitoid, Megaselia (M) scalaris (Diptera: Phoridae) of fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae) from India. Egypt. J. Biol. Pest Control 31: 1–4. https://doi.org/10.1186/s41938-021-00439-w.

Sharanabasappa, D., C.M. Kalleshwaraswamy, J. Poorani, M.S. Maruthi, H.B. Pavithra and J. Diraviam (2019). Natural enemies of Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae), a recent invasive pest on maize in South India. Fla. Entomol. 102(2): 619−623. https://doi.org/10.1653/024.102.0335.

Sharanabasappa, D., C.M. Kalleshwaraswamy, R. Asokan, H.M. Swamy, M.S. Maruthi, H.B. Pavithra and G. Goergen (2018). First report of the fall armyworm, Spodoptera fru­giperda (J E Smith) (Lepidoptera: Noctuidae), an alien inva­sive pest on maize in India. Pest Manag. Hort. Ecosyst. 24: 23–29.

Shylesha, A. N., S.K. Jalali, A. Gupta, R. Varshney, T. Venkatesan, P. Shetty, C.R. Ballal  (2018). Studies on new inva­sive pest Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) and its natural enemies. J. Biol. Control. 32(3) :145. https://doi.org/10.18311/jbc/2018/21707.

Singhamuni, A., A. Abdul Aziz and N. Ashikin Abdullah (2025). Life cycle and predation of Sycanus dichotomus (Hemiptera: Reduviidae) on Spodoptera  frugiperda larvae. The Indian J. Agric. Sci. 95(11): 1393-1399.https://doi.org/10.56093/ijas.v95i11.155179.

Sisay, B., J. Simiyu, E. Mendesil, P. Likhayo, G. Ayalew, S. Mohamed and T. Tefera  (2019b). Fall armyworm, Spodoptera frugiperda infestations in East Africa: Assessment of damage and parasitism. J. Insects 10(7): 195. https://doi.org/10.3390/insects10070195.

Sisay, B., J. Simiyu, P. Malusi, P. Likhayo, E. Mendesil, N. Elibariki, M. Wakgari, G. Ayalew and T. Tefera (2018). First report of the fall armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae), natural enemies from Africa. J. Appl. Entomol.  142:800–804. https://doi.org/10.1111/jen.12534.

Sivakumar, G., M. Kannan, S. Ramesh Babu, M. Mohan, M. Sampath Kumar, P. Raveendran T. Venkatesan, R. Rangeshwaran, C.R. Ballal and P. Ram Kumar (2020). Isolation and characterization of indigenous nucleopolyhedrovirus infecting fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) in India. J.Curr. Sci. 119 (5): 860-864. https://doi.org/10.18520/cs/v119/i5/854-860.

Sousa, E. H. S., M.C.B. Matos, R.S. Almeida and A.V. Teodoro (2011). Forest fragments’ contribution to the natural biological control of Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) in Maize. Braz. Arch. Biol. Technol. 54(4):755–760. https://doi.org/10.1590/S1516-89132011000400015.

Southon, R. J., O.A.  Fernandes, F.S. Nascimento and S. Sumner (2019). Social wasps are effective biocontrol agents of key lepidopteran crop pests. Proc. R. Soc. B 286(1914) :20191676. https://doi.org/10.1098/rspb.2019.1676.

Sujeetha, J.A.R.P. and K. Sahayaraj (2014). Role of Entomopathogenic Fungus in Pest Management. Springer: New Delhi (India). 31-46 p.

Sun, B., F. Li, X. He, F. Cao, E. Bandason, D. Shapiro-Ilan   and S. Wu (2020). First report of Ovomermis sinensis (Nematoda: Mermithidae) parasitizing fall armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) in China. J. Nematol. 52: 1. https://doi.org/10.21307/jofnem-2020-050.

Supeno, B., H. Haryanto, T. Tarmizi and N.M.L.T. Eranawati (2021). Parasitoid of fall armyworm larvae, Spodoptera frugiperda (Lepidoptera: Noctuidae) on maize at Lombok Island, 33, 00016, ICST conference, December 14th 2020, published online: June 1st 2021. 2: 460-466.

Tang, J., B. Lyu, H. Lu, X. JI, P. Yang, H. Su and B. Cai (2020a). Investigation and preliminary study of biological characteristic of parasitic wasps of Spodoptera frugiperda in Hainan. Redai Zuowu Xuebao, 41(6): 1189-1195. https://doi.org/ 0.3969/j.issn.1000‑2561.2020.06.017.

Tang, Y. T., Y.Y.  Li, C.X. Liu, J.J Mao, H.Y.  Chen, L.S. Zhang and M.Q. Zhang (2019). Predation and behavior of Arma chinensis (Fallou) to Spodoptera frugiperda (JE Smith). Plant Prot.45(4): 65-68. https://doi. org/10.16688/j.zwbh.2019264.

Tang, Y., J. Guo, Q. Wang, H. Tai, Z. Cao, K. He and Z. Wang (2020b). Three larval parasitic wasps of Spodoptera frugiperda were found in Dehong prefecture of Yunnan province. J. Plant Prot. Res. 46: 254–259.

Tang, Y., M. Wang, H. Chen, Y. Wang, H. Zhang, F. Chen and L. Zhang (2019). Predatory capacity and behavior of Picromerus lewisi Scott against Spodoptera frugiperda higher instar larve. Chin. J. Biol. Control. 35(5): 698-703. https://doi.org/ 10.16409/j.cnki.2095‑039x.2019.04.005.

Tang, Y., Q. Li, L. Xiang, R. Gu, Y. Wu, Y. Zhang and Z. Zhou (2021). First report on Megaselia scalaris Loew (Diptera: Phoridae) infestation of the invasive pest Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) in China. J. Insects, 12(1): 65. https://doi.org/ 10.3390/insects12010065.

Tawakkal, M. I., D. Buchori, N. Maryana and Pudjianto (2021). May. New association between Spodoptera frugiperda JE Smith (Lepidoptera: Noctuidae) and native natural enemies: Bioprospection of native natural enemies as biological con­trol agents. [IOP Publishing.]. IOP Conference Series. Earth Environ.  Sci. 771(1): 012030. https://doi.org/10.1088/1755-1315/771/1/012030.

Tendeng, E., B. Labou, M. Diatte, S. Djiba and K. Diarra (2019). The fall armyworm Spodoptera frugiperda (J. E. Smith), a new pest of maize in Africa: biology and first native natural enemies detected. Int. J. Biol. Chem. Sci. 13(2): 1011-1026. https:// doi.org/10.4314/ ijbcs.v13i2.35.

Valicente, F.H. (1988). Leaf consumption of the corn fall armyworm, Spodoptera frugiperda (J. E. Smith, 1797) infected with granulosis or nuclear polyhedrosis virus. An. Soc. Entomol. Brasil. 17(2): 347–357.

Van Lenteren, J.C. (2012). The state of commercial augmentative biological control: plenty of natural enemies, but a frustrating lack of uptake. Bio Control. 57(1): 1–20. https://doi.org/10.1007/s10526-011-9395-1.

Varella, A. C., A.C. Menezes-Netto, J.D.S. Alonso, D.F. Caixeta, R.K.D. Peterson and A.O. Fernandes, (2015). Mortality Dynamics of Spodoptera frugiperda (Lepidoptera: Noctuidae) Immatures in Maize. PLoS One, 10(6): e0130437. https://doi.org/10.1371/journal.pone.0130437.

Vercher, R., J. Costa-Comelles, C. Marzal and F. Garcia-Mari (2005). Recruitment of native parasitoid species by the invading leafminer Phyllocnistis citrella (Lepidoptera: Gracillariidae) on citrus in Spain. Environ. Entomol. 34(5): 1129–1138. https://doi.org/ 10.1093/ee/34.5.1129.

Wahyuningsih, R. D., T. Harjaka, Y. Suputa and A. Trisyano (2022). Parasitization levels of Spodoptera frugiperda eggs in three different corn ecosystems in East Java.J.  Perlin. Tanam, Indonesia,26 (1):28-39. https://doi.org/10.22146/jpti. 71598.

Wu, C., L. Zhang, C. Liao, K. WU, Y. Xiao and Y. Xiam (2019). Research progress of resistance mechanism and management techniques of fall armyworm Spodoptera frugiprerda to insecticides and Bt Crops. J. Plant Dis. Pests, 10 (4): 10-178.https://doi.org/10.19579/j.cnki.plant-d.p.2019.04.004.

Yang, J. G., M. Zhao, P. Zhu, Z.H. Wang and Y.N. Li (2019). Field experiment of release Trichogramma chilonis Ishii on Spodoptera frugiperda (Smith). China Plant Prot. 39: 59–61.

Yang, L., F. Li, X. Lu, B. Xing, X. Pan, X. Shi   and S. Wu (2022). Performance of three Trichogramma species as biocontrol agents on Spodoptera frugiperda eggs. J.  Appl. Entomol. 146(8): 1019-1027. https://doi.org/ 10.1111/jen.13042.

Yu, S. (1991). Insecticide resistance in the fall armyworm, Spodoptera frugiperda (J.E Smith). Pestic. Biochem. Physiol, 39: 84-91.https://doi.org/10.1016/0048-3575(91)90216-9.

Yu, S.,S. Nguyen and G. Abo-Elghar (2003). Biochemical characteristics of insecticide resistance in the fall armyworm. Spodoptera frugiperda (J.E Smith). Pest. Biochem. Physiol. 77: 1-11.https://doi.org /10.1016/S0048-3575(03)00079-8.

Zang, L. S., S. Wang, F. Zhang and N. Desneux (2021). Biological control with Trichogramma in China: History, present status, and perspectives. Annu. Rev. Entomol. 66(1): 463–484. https://doi.org/10.1146/annurev-ento-060120-091620.

Zeng, G., J.R. Zhi, C.R. Zhang, T. Zhang, J.Q. Ye, L. Zhou and M. Ye (2021). Orius similis (Hemiptera: Anthocoridae): A promising candidate predator of Spodoptera frugiperda (Lepidoptera: Noctuidae). J. Econ. Entomol. 114(2): 582-589. https://doi.org/ 10.1093/jee/toaa318.

Zhao, X., K.H. Zhu, Z.T. Zhang, K.L.  He, L.S.  Zhang, J.C. Zhou and H. Dong (2020). Preliminary evaluation of the control efficacy of Telenomus remus against Spodoptera frugiperda in field conditions. Plant Prot. 46: 74–77. https://doi.org/ 10.16688/j.zwbh.2019533.