Spodoptera frugiperda adults are highly active at night, seeking mates and expanding into new territories while staying elusive during the day (Patel Sagarbhai et al., 2021). Research has confirmed that the males of FAW are more attracted to light than females (Vilarinho et al., 2011). Studies have also documented that FAW moths from the same generation can travel more than 500 kilometers before they are ready to lay their eggs (Prasanna et al., 2018; Acharya et al., 2020; Badhai et al., 2020). Male and female S. frugiperda are shown in Figures 1a, b , respectively.

Oviposition typically occurs at night, with females laying inconspicuous egg clusters on the undersides of leaves to shield them from predators (Kasige et al., 2022). In some cases, the eggs are laid in layers and covered with scales for added protection. The upper leaf surface and whorl of host plants may also harbor some egg batches (Sharanabasappa et al., 2018). A single FAW female can lay an average of 1500 eggs, showing the high fecundity of the pest (Prasanna et al., 2018). FAW eggs hatch into tiny neonate larvae within 3 to 4 days at an average temperature of 28°C. Eggs of S. frugiperda are shown in Figure 1c .

Soon after hatching, the larvae disperse from the site and infest nearby plants, often aided by the wind (Ortiz-Carreón et al., 2024). The larval stage of FAW consists of six instars and lasts about 14 days in warm conditions and up to 30 days in cooler weather (Prasanna et al., 2018). This stage is the damaging phase of FAW’s life cycle (Sagar et al., 2020). Caterpillars primarily feed at night, but activity may also occur in the late evening or early morning (Schlemmer, 2018; Patel Sagarbhai et al., 2021). They feed on the leaves, the tender stem, the silks, and the ears of host plants. As they grow, their appetite intensifies, causing severe damage to host plants. Day et al. (2017) and Flanders et al. (2017) reported that the largest larvae can cause up to 77% of plant damage.

During the vegetative phase of the host crops, continuous feeding results in skeletonized leaves and heavily windowed whorls filled with larval frass (Goergen et al., 2016).

Cannibalism is an important biological phenomenon within the FAW larval stage. This intraspecific behavior frequently results in a significant reduction in larval numbers, leaving only one larva per plant. The eliminated cohabitant either succumbs to cannibalism or relocates to another plant where competition is less intense. According to Ren et al. (2020) and Ahissou et al. (2022), a high density of larvae on a maize plant suggests that the larvae have not yet reached advanced developmental stages. Chapman et al. (2000) observed that the larvae of FAW exhibit cannibalistic behavior from the third instar (L3) stage onward, allowing them to dominate interspecific competitors and reduce intraspecific competition. Larva of S. frugiperda is shown in Figure 1d .

At the 6^th instar, larvae undergo metamorphosis within a woven silky cocoon. Pupation usually occurs in the soil ( Figure 1e ) at a depth of 2-8 cm (Ojumoola, 2021). During summer, the pupal stage lasts about 8-9 days. Studies conducted by Huang et al. (2021) and Montezano et al. (2018) revealed that the pupal developmental stage was shorter in female FAW than in males. After pupation, adult moths emerge and live for an average of 14 days (Prasanna et al., 2018). The total life cycle of FAW (a generation time) is about 30–40 days at 28°C but extends to approximately 55 days in cooler seasons (Prasanna et al., 2018; Ashok et al., 2020; Lekha et al., 2020). This is supported by Kenis et al. (2022), who found that the FAW life cycle can extend to 90 days at lower temperatures. Tendeng et al. (2019) found that the total cycle is between 22 and 28 days at 25°C with an average of 25 days in laboratory conditions.

Environmental factors such as temperature, relative humidity, soil moisture, and host plant characteristics (quality, diversity, availability and phenology), and the presence of natural enemies influence the development, distribution, abundance and population dynamics of FAW (Kenis et al., 2022).

Several studies have demonstrated the direct impact of temperature and relative humidity on various biological activities, including adult mating, egg production and hatching, larval development, pupal survival, and adult longevity ( Table 1 ). Studies have indicated that FAW egg-to-adult development requires a specific number of degree days, with potential differences in developmental duration between sexes (Schlemmer, 2018; Malekera et al., 2022). Additionally, research has shown that larval and pupal development is optimal at temperatures ranging from 28 to 30°C, while lower temperatures tend to prolong the developmental period compared to warmer temperatures (Du Plessis et al., 2020). Schlemmer (2018) and Kumara et al. (2022) noted that temperatures around 30°C favor the growth and development of FAW, resulting in the fastest larval growth rate and the lowest mortality. Gergs and Baden (2021) found that lower ambient temperatures, combined with poor diet quality, increase the duration of FAW’s life cycle.

Researchers typically observed the effect of temperature on pupal weight, noting the highest weights at 25°C and decreasing weights as temperatures deviate from this optimal point (Huang et al., 2021).

Rising temperatures due to global warming may increase the number of FAW generations, facilitating its spread and establishment in new regions (Garcia et al., 2018; Yan et al., 2022). In African climatic conditions, the pest reproduces continuously, producing up to 15 generations annually (Tendeng et al., 2019; Barkessa et al., 2024).

Studies have shown that relative humidity (RH) also plays an important role in FAW biology. According to He et al. (2021), an RH of 80% was optimal for FAW to achieve its highest intrinsic rate of increase, finite rate of increase, and net reproduction rate. Additionally, the authors noted that soil moisture ranging from 6.80% to 47.59% supports FAW pupation, survival, and eclosion.

Another key factor influencing the ecology and population dynamics of FAW is host plant selection. The preference of FAW for a particular host plant depends on the strain type (Tiwari, 2022). Two distinct strains have been identified within the FAW population: the C-strain and R-strain (Schofl et al., 2009; Groot et al., 2010; Dumas et al., 2015; Jing et al., 2021). Although these strains have morphological similarities, they differ in several key biological aspects including host plant preference, reproductive behaviors, genetic profiles, and pheromone compositions. The C-strain predominantly feeds on maize, sorghum, and cotton, whereas the R-strain demonstrates a greater affinity for rice, sugarcane, and pasture grasses (Dumas et al., 2015; Jing et al., 2021).

Beyond host specificity, these strains also exhibit differences in their dispersal abilities and resistance to environmental stressors. Studies suggest that the C-strain is more adapted to agricultural landscapes with annual crops, while the R-strain thrives in more stable, perennial grassland environments (Nagoshi et al., 2017). Additionally, inter-strain hybridization has been reported, potentially leading to increased genetic diversity and adaptive capabilities in invasive populations (Nagoshi et al., 2019). The presence of both strains in Africa has raised concerns about their combined impact on food security, as they contribute to the rapid expansion and persistence of FAW populations across diverse agroecosystems (Assefa, 2019).

Regular scouting of crops for signs of FAW infestations can help with early detection and prompt action before populations reach the economic threshold levels. According to Prasanna et al. (2018), implementing effective monitoring, surveillance, and scouting systems is a critical step in developing a successful IPM of FAW. Essential tools and practices for field monitoring at the farmer level include regular field scouting, use of pheromones (Tepa-Yotto et al., 2022b), and light traps. Pheromone traps are one of the most prominent and effective scientific tools used in pest management programs worldwide. This device enables early detection of pest infestations, tracks pest population dynamics, and supports pest management decision-making (Chiwamba et al., 2018). For small-scale farmers, several advanced technologies are available for monitoring and surveillance of FAW.

The FAW Monitoring and Early Warning System (FAMEWS) has been widely adopted across the continent, enabling farmers to track pest infestations (FAO, 2020). In six Eastern African countries (Ethiopia, Kenya, Tanzania, Uganda, Rwanda, and Burundi), a community-based initiative involving 650 focal persons has demonstrated its effectiveness (Niassy et al., 2021).

Remote sensing, particularly satellite imagery and NDVI analysis, has proven useful for detecting FAW damage. A Sentinel-2 satellite-based algorithm successfully identified biomass loss in maize fields in Zimbabwe, Kenya, and Tanzania (Buchaillot et al., 2022). In Ghana, integrating field surveys with remote sensing improved FAW distribution modeling (Bilintoh, 2019). Predictive models, such as MaxEnt, have been used to simulate FAW distribution under current and future climate scenarios in Ethiopia (Barkessa et al., 2024).

Machine learning is also being explored, with the University of Zambia developing an automated FAW identification system using Convolutional Neural Networks (CNN) that integrates vision and motion sensors with pheromone traps for real-time pest monitoring, reducing field visits, improving data collection efficiency, and enhancing decision-making (Chiwamba et al., 2018).

Studies have found that cultural practices such as crop rotation, regular weeding, and destruction of crop residues help to mitigate FAW infestation (Cokola et al., 2023). Maintaining organic soil fertility was found to prevent high FAW density (Bayissa et al., 2023). Using early-maturing cultivars was reported to be effective against FAW, as these varieties have a shorter growth cycle, reducing their exposure to the pest (Day et al., 2017). Also, adjusting the planting schedule to avoid peak periods of FAW activity can help prevent infestations (Ahissou et al., 2022). Other studies have indicated that destroying the ratoons of sorghum and sugarcane, which serve as reservoirs for FAW, also effectively controls the pest’s population. Intercropping maize or other cereals with non-preferred or repellent crops (e.g., legume crops, desmodium and Dolichos lablab), as well as practicing mixed cropping, has been shown to attract natural enemies that prey on FAW, making it a proven effective pest management strategy (Ratnadass et al., 2006; Chimweta et al., 2020). However, many leguminous crops that are often grown with maize are known to be hosts for FAW and tend to sprout earlier, which could exacerbate the pest infestations in maize (Rwomushana et al., 2018; Ambia, 2023). Other practices like zero-tillage can reduce overwintering larvae and promote higher densities of natural enemies, making the environment less attractive for FAW populations (Baudron et al., 2019). The number of FAW predators can also be maintained through habitat management techniques like building shelterbelts or live fences that increase biodiversity (Hellmich et al., 2008). Additionally, push-pull technology, which combines trap plants to attract FAW and repellent intercrops to deter it, is cited as an ideal approach for mixed farming systems, effectively reducing FAW larval density and damage to maize and other cereals (Joatya et al., 2022; ICIPE, 2024). However, certain cultural practices, like intercropping cereals with pumpkins, should be avoided as they may inadvertently increase FAW damage by providing shelter for moths and facilitating larval migration. Integrating these cultural practices with other FAW control strategies can significantly improve the management of the pest.

Host plant resistance (HPR) is a key component of IPM strategies for controlling FAW. It involves the inherent defense mechanisms of plants, which include both biophysical and biochemical properties (Meihls et al., 2012; Jin, 2017);.

Studies discovered several traits associated with maize resistance against FAW. Callahan et al. (1992) found that a combination of two specific polypeptides (36 and 21 kDa) is predictive of resistance to lepidopteran insects. Furthermore, Smith et al. (2012) revealed that the terpenoid (E)-β-caryophyllene in maize whorl leaves showed that the plant is naturally resistant to FAW. Other compounds associated with resistance include mir1-CP, oxophytodienoic acid (OPDA), a low protein/total nonstructural carbohydrates ratio, thicker leaves, and higher contents of crude fiber, hemicellulose, and thicker cell wall complexes, as well as the constitutive expression of jasmonic acid genes (Singh et al., 2022). Similarly, Darshan et al. (2024) reported significant negative correlations among morphological traits such as plant height, stem girth, and the presence of trichomes related to FAW incidence. Baudron et al. (2019) and Snook et al. (1993) stated that the rapid accumulation of proteins or phytochemicals like maysin in silks can either poison or starve the pest. Chiriboga Morales et al. (2021) and dos Santos et al. (2020) found a negative correlation between maize leaf toughness at the twelfth vegetative leaf stage (V12) and leaf damage from FAW infestation. Jiang et al. (1995) and Pechan et al. (2000) found that higher levels of the 33-kD cysteine proteinase in plant leaves were associated with lower FAW larval weight, suggesting that the enzyme may serve as a novel insect defense mechanism in plants. Baudron et al. (2019) reported that plant characteristics such as the density of leaf hairs or the density of the cuticular wax layer can reduce foliar damage. In response to pest attacks, some plants activate wound-response pathways and release volatile compounds (Smith et al., 2012).

To date, the International Maize and Wheat Improvement Center (CIMMYT) has disseminated several FAW-tolerant maize inbred lines, including CML71, CML124, CML125, CML338, CML333, CML334, CML370, CML372, and CML574, across 34 countries. Also, the center has developed three high yielding maize hybrids (FAWTH2001, FAWTH2002 and FAWTH2003) with native tolerance to FAW, which are being widely released in Africa. CIMMYT undertook these research efforts in collaboration with various National Agricultural Research and Extension Systems (NARES), advanced research institutes (ARIs), and commercial seed companies (Prasanna et al., 2021). Several other studies have identified different maize germplasms with desirable resistance traits or mechanisms to FAW, as illustrated in Table 2 .

The identification of resistant donor lines and the introduction of desired traits into elite maize cultivars represent significant advancements in developing FAW-resistant plants. Exploring molecular markers for FAW resistance can accelerate the breeding process (Sharma et al., 2008; Nashath et al., 2023). Two important tools used in modern breeding are marker-assisted selection and quantitative trait loci (QTLs). They facilitate the identification of genetic traits, accelerate breeding cycles, and enhance the development of insect-resistant germplasms (Mihm, 1997; Singh et al., 2022). These practices help incorporate desired traits while avoiding deleterious alleles. According to Abdulmalik et al. (2017), molecular tools make it possible to select for FAW resistance while keeping the beneficial traits of adapted cultivars. Singh et al. (2022) emphasized that rapid screening strategies and genomic selection can enhance the utilization of plant genetic resources. In a GWAS study, Badji et al. (2020) utilized SNP markers to identify 62 quantitative trait nucleotides (QTNs) linked to FAW resistance traits in maize in four SNP regions. Notably, QTLs on chromosomes 4 and 9 correspond to bi-parental QTL mapping, highlighting their importance they in in developing markers for FAW leaf damage resistance (Womack et al., 2018; 2020).

Emerging biotechnological solutions are being explored to combat lepidopteran pests, including stemborers and FAW. In the African context, genetically modified Bt maize, which is resistant to both pests, has been commercialized in South Africa and is expected to expand to other countries where transgenic maize is already approved as an effective management strategy (Prasanna et al., 2022). Furthermore, advanced molecular technologies, including RNA interference (RNAi) (Kebede and Fites, 2022; Kumara et al., 2022; Nwokeoji et al., 2022) and gene editing using CRISPR-Cas9 (Singh et al., 2022), have been investigated to target essential genes in lepidopteran pests, disrupting their development and reducing crop damage. However, these technologies have yet to be implemented for FAW management in Africa.

It has been reported that practices like handpicking larvae, crushing egg and neonate masses, and uprooting infested plants increase the resilience to FAW. However, these options are best suited for shorter cultivars, early to mid-whorl stages, and small-sized fields (Chimweta et al., 2020). Traditionally, small-scale farmers in Africa apply soil and ash to leaf whorls to control the pest (Abate et al., 2000; Ambia, 2023). These mechanical methods are still applicable at the farmer level in SSA due to the relatively small areas of production (Hruska, 2019; Cokola et al., 2023). Other mechanical approaches used in SSA to control FAW include the use of pitfall traps, sticky traps, light traps, or pheromone traps (Prasanna et al., 2018; Bhusal and Bhattarai, 2019; Gebreziher and Gebreziher, 2020; Akeme et al., 2021), as well as ash and detergents (Tambo et al., 2020). Also, legumes such as beans (Phaseolus vulgaris), groundnuts (Arachis hypogaea), cowpeas (Vigna unguiculata) and soybean (Glycine max) are intercropped within maize to serve as trap crops for the pest (Akeme et al., 2021).

Biological control is an effective strategy for managing FAW using natural enemies such as predators, parasitoids, entomopathogenic fungi, viruses, nematodes, bacteria, and biopesticides (Assefa and Ayalew, 2019; Abbas et al., 2022). This can be implemented through two primary approaches: augmentative biocontrol, which involves mass rearing and periodic release of beneficial organisms to boost existing populations, and inoculative biocontrol, which entails a single initial release of natural enemies aimed at establishing a self-sustaining population for long-term pest control (Nyamutukwa et al., 2022).

Additionally, proper habitat conservation, maintaining prey availability, and minimizing the use of broad-spectrum insecticides in agricultural and natural ecosystems can support the populations of natural enemies, a practice known as conservation biocontrol. These methods can be effectively integrated as biocontrol-based applications within an IPM framework against FAW (Tefera et al., 2019; Kenis et al., 2022; Ayra-Pardo et al., 2024).

Ideally, biological control should be based on natural enemies already attacking FAW in Africa rather than organisms from other regions, as these may cause unforeseen problems (Chimweta et al., 2020). In Africa, Abbas (2023) reported 48 species of parasitoids, primarily from the Braconidae family (19 species), followed by Ichneumonidae (11 species), Tachinidae (9 species), and Trichogrammatidae (5 species). The remaining species belong to families such as Eulophidae, Heciridae, Pteromalidae, and Scelionidae, with one species each. The egg-larval endoparasitoid Chelonus bifoveolatus (Hymenoptera: Braconidae) commonly targets the eggs of FAW on the African continent, causing the young instars of FAW to hatch alongside the parasitoid’s eggs (Obala et al., 2023; ICIPE, 2024). In Africa, Trichogramma chilonis (Hymenoptera: Trichogrammatidae), Coccygidium luteum, Telenomus remus (Hymenoptera: Scelionidae), and Cotesia icipe are also common FAW parasitoids. Cotesia icipe is known for its strong ability to control FAW in maize crops.

Several studies found that the parasitism rates of those mentioned parasitoids varied across African countries, such as Ghana, Kenya, Niger, Nigeria, Egypt, Uganda, South Africa, Senegal, Tanzania, and Zambia (Agboyi et al., 2020; Abang et al., 2021; Mohamed et al., 2021; Otim et al., 2021; Abbas, 2023; Koffi et al., 2023; ICIPE, 2024).

Trichogramma spp. and Telenomus spp. are widely used parasitoids for FAW control because they are easy to rear in laboratories and effective in managing FAW population (Figueiredo et al., 2002; Gutierrez-Martinez et al., 2012; Tefera et al., 2019).

Abbas (2023) reported a total of 17 predators of FAW in Africa, which include seven coleopterans from the Coccinellidae family (Coleomegilla maculata, Cycloreda sanguinea, Cheilomenes sulphurea, Coccinella transvirsalis, Harmonia octomaculata, Eriopis sp., and Hippodamia sp.), two coleopterans from the Carabidae family (Callida sp. and Calosoma granulatus), one coleopteran from the Lampyridae family (Hematochares obscuripennis), three dermapterans from the Forficulidae family (Diaperasticus crythrocephalus, Forficula senegalensis, and Doru sp.), one hemipteran from the Geocoridae family (Geocoris sp.), one hemipteran from the Pentatomidae family (Podisus sp.), one hymenopteran from the Formicidae family (Pheidole megacephala), and one lepidopteran from the Erebidae family (Perprius modiulipes). Table 3 shows the main families and order/class of FAW’s natural enemies in Africa.

Several bio-rational control agents have shown effectiveness in FAW management. These include the entomopathogenic fungi Metarhizium anisopliae and Beauveria bassiana, as well as the biological pesticide Spodoptera frugiperda nucleo-polyhedrovirus (Fawligen, SfNPV) (Akutse et al., 2019). Nematodes, protozoans, and botanical extracts such as powders, volatile and non-volatile oils, and extracts from Azadirachta indica and Jatropha curcas seed can also be used to control FAW (Adjaoke et al., 2021; Abbas et al., 2022; El-Sappagh et al., 2022; Tepa-Yotto et al., 2022b). Patel Sagarbhai et al. (2021) stated that neem affects insect growth, development, egg laying, and has been shown to have antifeeding and larvicidal activity in FAW. Sisay et al. (2019) noted that extracts from several plants, such as Milletia ferruginea, Phytolacca dodecandra, Schinus molle, Melia abyssinica, Nicotiana tabacum, Lantana camara, Chenopodium ambroides, and Jatropha gossypifolia, induced high mortality of FAW. The deployment of biological control methods against FAW among African farmers has been progressing. However, adoption remains minimal due to the absence of true native natural enemies. In Zambia, CABI launched a village-based biological control initiative to promote the use of natural enemies and biopesticides, aiming to reduce farmers’ reliance on chemical pesticides (CABI, 2023). Additionally, training programs have been conducted to empower young farmers to explore business opportunities in the biocontrol sector, further facilitating the adoption of these methods (CABI, 2024).

At present, most African farmers rely on the application of synthetic insecticides to control the FAW (Assefa and Ayalew, 2019). According to Tambo et al. (2020) and Kumela et al. (2019), the most used control measure in Africa for the management of FAW is the application of synthetic pesticides.

Insecticide applications should be based on FAW infestation thresholds after continuous monitoring. Studies have validated several treatment thresholds that consider not only the pest’s developmental stages but also the phenological stages of the infested crops (FAO, 2017; Prasanna et al., 2018). To be technically and cost-effective, the pesticide choice should respect these action thresholds (Chimweta et al., 2020).

However, chemical insecticides should be considered as the last resort for managing FAW, as they may negatively impact the environment and the non-target organisms, human health, and biodiversity, including beneficial insects (Harrison et al., 2019; Deshmukh et al., 2020; Idrees et al., 2022). Furthermore, their use is associated with high costs, potential environmental contamination, resistance development, and frequent pest resurgences (Gebreziher, 2020). Studies have demonstrated that repeated, indiscriminate use and misuse of chemical insecticides exert strong selection pressure, leading to the evolution of resistance mechanisms (Togola et al., 2018; Zhang et al., 2021). Cases of FAW resistance to multiple classes of insecticides, including pyrethroids, organophosphates, and carbamates, have been widely documented in different regions, posing a significant challenge to pest management (Carvalho et al., 2013; Yu et al., 2023). The increasing resistance not only reduces the efficacy of chemical control but also requires higher application rates and more frequent spraying, further exacerbating environmental and economic burdens (Gutiérrez-Moreno et al., 2019). To prevent insecticide resistance in FAW, it is important to implement integrated pest management practices. These strategies will help maintain resistance at low levels.

Studies have shown that selection pressure from Bacillus thuringiensis (Bt)-resistant maize can lead to FAW resistance. FAW has developed resistance to Bt toxins such as Cry1Ac, Cry1Ab, and Cry1A.105 through mechanisms like metabolic detoxification, target-site insensitivity, and behavioral avoidance (Storer et al., 2010; Huang et al., 2014; Boaventura et al., 2020; Boaventura et al., 2021; Jing et al., 2021; Zhang et al., 2021). This poses significant challenges for FAW- preferred crop growers (Peferoen, 1997; Zhang et al., 2021). Several studies have also shown that FAW has become resistant to different ranges of pesticides, such as carbamates, organophosphates, and pyrethroids (Jing et al., 2021; Zhang et al., 2021).

It is suggested that to break insects from becoming resistant to protein toxins, an insect resistance management (IRM) approach is recommended. One key strategy of IRM is the establishment of refuge areas, either within or adjacent to transgenic fields, to allow non-resistant insect populations to thrive. This promotes random mating between treated and refuge populations, weakening the resistance alleles in the overall population (Vilarinho et al., 2011; Murúa et al., 2019). Several studies have shown that the presence of non-Bt refuges can sustain the long-term efficacy of Bt crops by preventing resistant populations from becoming dominant (Huang et al., 2014; Carrière et al., 2020). Proper implementation of refuge areas requires careful planning, including the selection of appropriate non-Bt host plants, strategic placement, and maintaining a sufficient proportion of the crop area as a refuge. Research suggests that an ideal refuge should be at least 20% of the total crop area (Bourguet et al., 2005; EuropaBio, 2019). Other studies suggest placing refuge areas approximately 800 meters apart to maintain their effectiveness (Vilarinho et al., 2011). Prasanna et al. (2022) suggested that integrating polygenic native resistance with transgenic Bt maize, which typically relies on one gene (monogenic) or a few genes (oligogenic) in conventional cropping systems, could be a key strategy for insect resistance management. This approach would help prevent FAW from overcoming the limited genetic defenses of Bt maize, thereby promoting a more effective and sustainable FAW control strategy.