OVERVIEW OF RICE LEAF FOLDER, Cnaphalocrocis medinalis AND UNDERSTANDING ITS MANAGEMENT THROUGH BIOLOGICAL, CHEMICAL AND GENETIC APPROACHES

INTRODUCTION

 Rice is one of the world’s most important cereal crops and a staple food for nearly half of the global population. It plays a critical role in food security, employment, economic growth, social stability worldwide (Maclean et al., 2013). Major rice-producing countries include India, China, Bangladesh, Indonesia , Vietnam, Thailand, Philippines, Burma, Pakistan, and Brazil, with annual production ranging from 8.68 to 150 million tons USDA Production Data (https://www.fas.usda.gov/data/production/commodity/0422110) (Figure 1). Globally, rice provides more than 20% of dietary energy and supplies over 70% of daily caloric intake in several Asian countries (Yadav and Kumar, 2018). Numerous insect pests attack rice at different growth stages. Among them, the rice leaf folder, Cnaphalocrocis medinalis (Guenée) (Lepidoptera: Pyralidae), is one of the most destructive pest of rice. Larvae fold rice leaves and feed on mesophyll tissues within the rolled leaves, causing significant annual yield losses estimated at 63-80% worldwide (Horgan, 2017). Management of C. medinalis has traditionally relied on biological control, cultural practices, and particularly synthetic insecticides (Su et al., 2003). However, effective control strategies face multiple challenges, including the pest’s high reproductive capacity, migratory behavior, insecticide resistance; global climate changes induced alterations in its distribution, development, and reproduction (Bodlah et al., 2017; Wang et al., 2017; Hajjar et al., 2023). This review summarizes relevant literature on the bioecology, geographical distribution, natural enemies, and management strategies and identifies future research priorities to address knowledge gaps and optimize integrated pest management strategies for C. medinalis.

Figure 1. Percentage of rice production by country in 2025

  A comprehensive literature search was conducted to collect and synthesize published information on the biology, ecology, host–pest interactions, monitoring, resistance mechanisms, and integrated management strategies of C. medinalis. The literature search was performed using reputable scientific databases, including Web of Science (WoS), Scopus, PubMed, Google Scholar, ScienceDirect, and SpringerLink. The search strategy employed combinations of the following keywords: “Cnaphalocrocis medinalis,” “rice leaf folder,” “rice leaf folder,” “biology,” “ecology,” “population dynamics,” “migration,” “host plant resistance,” “integrated pest management,” “biological control,” “entomopathogenic fungi,” “entomopathogenic nematodes,” “RNA interference,” “Bacillus thuringiensis,” “insecticide resistance,” “gut microbiota,” “monitoring,” and “climate change.” Publications from 1959 to 2025 were considered, search was restricted to peer-reviewed journal articles, books, book chapters, conference proceedings, and technical reports published in English. Approximately 450 publications were initially identified from all databases, with the majority retrieved from Web of Science and Scopus, while additional records were obtained from Google Scholar, PubMed, and reference lists of relevant publications. After removing duplicate records, the titles and abstracts were screened for relevance. Full-text articles were then evaluated according to predefined inclusion criteria, including studies focusing on the taxonomy, biology, ecology, behavior, migration, host plant interactions, insecticide resistance, molecular biology, genomics, RNA interference, biological control, ecological engineering, and integrated pest management of C. medinalis. Publications lacking sufficient methodological details, unrelated studies, duplicate records, non-peer-reviewed articles, and publications without accessible full texts were excluded. Following the screening and eligibility assessment, 220 publications were selected and included in this review to provide a comprehensive and up-to-date synthesis of current knowledge on C. medinalis and its sustainable management.

Taxonomy and morphology:  C. medinalis was first described by Achille Guenée in 1854 and was previously reported under the synonyms Salbia medinalis Guenée and Botys nurscialis Walker (Walker, 1859). The genus Cnaphalocrocis comprises approximately 36 species distributed mainly across Asia, of which nearly eight species infest gramineous crops by folding or rolling leaves (Mashhoor et al., 2019). However, only Marasmia patnalis, M. exigua, and C. medinalis are considered economically important pest (Liao et al., 2017). Due to morphological similarities and comparable feeding habits, C. medinalis is often misidentified with M. exigua at different developmental stages (Barrion et al., 1991). Nevertheless, adults can be distinguished based on coloration, body size, and wing patterns (Rani et al., 2007). Larvae can be differentiated by coloration, size, subdorsal prothoracic spots, pronotum characteristics, and the presence of inverted M shaped markings on the head capsule (Rani et al., 2007) (Figure 2).

Other morphological characteristics of C. medinalis are summarized below;

Egg: Eggs are oval, flattened, and approximately 1 mm long and 0.5 mm wide, with fine reticulations on the chorion surface. They are laid singly or in small groups, usually inside folded leaves (Pathak and Khan, 1994).

Larva: Larvae possess a light-brown head and a light yellow to green abdomen with distinct subdorsal spots on the prothoracic region. Mature larvae is 14-19 mm in length and become orange-red before pupation (Heong, 1993).

Pupa: Pupae are cylindrical, 7-10 mm long, and pointed at the posterior end. Newly formed pupae are light yellow but gradually turn reddish-brown to dark brown, with a darker dorsal region and lighter abdomen (Heong, 1993).

Adult: Adult moths are 7-9 mm long with a wingspan of 12-18 mm. The body and wings are yellowish brown and the wings are held obliquely while resting. The forewings possess three dark-brown transverse lines, of which the middle line is shorter and thicker. The hindwings contain two transverse lines; the inner line is short and does not extend to the trailing edge, whereas the outer line extends completely to the trailing edge (Dale, 1994).

Figure 2. Morphology of C. medinalis

Life cycle and biology: The optimum temperature required for egg production is 25-26 °C (Lv et al., 2021). A female lays approximately 50-300 eggs singly or in groups of 3-8 along the midrib of rice leaves (Pathak and Khan, 1994). Eggs hatch within about 5 days after oviposition. Newly emerged larvae are creamy white and later become yellow to yellowish green with a brownish head capsule. Neonate larvae feed gregariously on chlorophyll within unopened leaves (Fraenkel et al., 1981). From the second instar onwards, larvae fold leaf blades using silken threads and feed internally by scraping the chlorophyll contents, producing characteristic longitudinal white streaks. In some cases, 2-3 leaves may be stitched together to form larger feeding shelters. Larval development is completed within 14–25 days through five instars (Gangwar, 2015). Prior to pupation, larvae become pinkish white and construct silken cocoons inside folded leaves by cutting and stitching leaf margins together and starts pupation (Elanchezhyan and Balakrishnan, 2020). Prior to pupation, larvae become pinkish white and construct silken cocoons inside folded leaves by cutting and stitching leaf margins together (Elanchezhyan and Balakrishnan, 2020). Pupation occurs within a single folded leaf. Newly formed pupae are light brown and gradually turn reddish brown within 6-10 days (Figure 3). Adults mostly emerge within an evening period (Heong, 1993). Oviposition generally begins 2-3 nights after mating and occurs mainly between dusk and midnight. Adults usually emerge during the evening. The pre-oviposition and oviposition periods last approximately 3 and 4 days, respectively. Adult females survive for 5-17 days, and the complete life cycle is completed within 23-40 days (Dale, 1994).

Figure 3. The life cycle of C. medinalis

Damage and economic impact: C. medinalis damages the rice plants exclusively during the larval stage (Liu et al., 2021). Continuous feeding removes chlorophyll, causing leaves to become whitish, papery, and eventually desiccated (Tiwari et al., 2021). Larval feeding reduces photosynthetic activity, weakens plant vigor, and may result in 20-30% grains yield loss (Litsinger et al., 2005). Although damage during the vegetative stage is often recoverable, infestations during the reproductive stage severely reduce the grain filling and yield. Under heavy infestations, extensive leaf destruction may completely devastate rice fields, resulting in substantial economic losses (Padmavathi et al., 2013). Due to its severe impact, C. medinalis was included in the Class I Crop Diseases and Insect Pests list by the Ministry of Agriculture and Rural Affairs of China in 2023. Between 2013 and 2022, this pest affected an average of 14 million hectares annually in China, causing estimated yearly grain losses of 350,700 t, while mitigated losses reached 3.67 million t (Fuyan et al., 2024).

Distribution and ecology: C. medinalis is widely distributed in tropical and subtropical rice-growing areas, active year-round, with peak activity typically occurring during the rainy season (Ali et al., 2019). In colder regions, this pest is primarily active from May to October, completing approximately 4-5 generations per year (Junaid and Khan, 2023). C. medinalis is commonly found in dark, over-fertilized rice paddies, multiple rice cultivation, increased irrigated areas, and pesticide overuse have all led to population C. medinalis (Roseli et al., 2021). Climatic factors, particularly temperature, humidity and rainfall significantly influence the occurrence, development, behavior, distribution, and population size of the C. medinalis (Morshed et al., 2020; Morshed et al., 2023). Historically, C. medinalis was considered a minor pest before the 1920s; however, its economic importance gradually increased in the late 1980s due to the intensification of rice cultivation and modern agricultural practices (Ranasinghe, 1992). The first serious outbreaks were reported in India during (Litsinger, 1994), followed by widespread outbreaks in China, Japan, and Korea between 1967 and 1981 (Heinrichs, 1985; Lee, 1992). Migration is an important ecological adaptation of C. medinalis and contributes significantly to its seasonal distribution and outbreak dynamics. Annual migration generally occurs from tropical to temperate rice-growing regions, with adults initiating migratory flights mainly during the evening (Chatterjee et al., 2023). Migrating adults typically fly at altitudes below 500 m and are capable of sustaining flight for up to nine consecutive nights (Wang et al., 2010). Unlike many migratory insects, mating and feeding do not significantly reduce migratory ability, and females can continue ovarian development and egg maturation during migration (Chang et al., 1980). Trap survey conducted between period 2003 and 2013 revealed migration periods ranging from 72 to 122 days, with a higher proportion of mated females captured during June and July, indicating that migration is not constrained by oogenesis-flight syndrome (Fu et al., 2014). Based on cumulative flight duration, C. medinalis populations are categorized as residence (AFD < 40 min), migratory (40–130 min min), and strong migratory (AFD>130 min), with average cumulative flight durations of 11, 82, and 232 min, respectively (Chapman et al., 2010; Zhang et al., 2015). In addition, C. medinalis exhibits strong remigration ability, with more than 90% of individuals capable of a first remigration and over 70% capable of a second remigration (Guo et al., 2019).

Insect rearing: C. medinalis is polyphagous and have several gramineous hosts, including millet, sorghum, wheat, maize, oats, sugarcane, and other wild grasses (Gao et al., 2023).  Extensive studies have been conducted on laboratory rearing of C. medinalis using rice seedlings, corn seedlings, and artificial diets (Waldbauer and Marciano, 1981). Early studies demonstrated that neonate larvae perform best when initially reared on rice seedlings before transfer to artificial diet (Shono and Hirano, 1989; Baik, 2006). Subsequent research optimized artificial diets supplemented with rice or corn leaf powder and essential nutrients to support larval growth and development (Guo et al., 2013; Xu et al., 2013). Optimized diets containing wheat germ  (8 g), rice leaf powder (3 g), corn powder (4 g), yeast (4 g), and casein (4 g) enabled completion of larval and pupal development within approximately 27 days (Wang et al., 2013). Later formulations further improved diet composition by incorporating distilled water (100 g), agar (1.50 g), wheat germs (6 g), soybean powder (8 g), casein (2 g), methylparaben (0.10 g), sorbic acid (0.10 g), yeast (4 g), natamycin (0.03 g), antibiotics (0.03 g), ascorbic acid (0.10 g), and vitamin mix (0.10 g) reducing the developmental period to approximately 21 days (Wang et al., 2014).

Transcriptomic, genomic, and microbiome Studies: C. medinalis has been extensively investigated for its toxicology, ecology, and physiology insecticide resistance, population dynamics, and migration behavior (Alvi et al., 2003). However, molecular studies on this pest have expanded only recently. The transcriptomic analysis generated over 23 million sequencing reads assembled into 44,941 unigenes of which 56.8% matched known proteins in the NCBI database (Li et al., 2012). Subsequent de novo transcriptome sequencing of C. medinalis generated 29,367,797 reads assembled into 63,174 unigenes with an average length of 753 bp. Among these, 31,810 unigenes were annotated against the NCBI Nr database and 10,043 were mapped to 285 KEGG pathways. The analysis identified 360 genes potentially associated with insecticide resistance and 16 genes involved in chitin metabolism of C. medinalis (Yu et al., 2015). Later, transcriptome sequencing using Illumina HiSeqTM2000/MiSeq technology from antennae, protarsus, and reproductive organs identified 102 chemoreception-related genes, including 29 odorant receptors (ORs), 15 ionotropic receptors (IRs), 30 odorant-binding proteins (OBPs), 26 chemosensory proteins (CSPs), and 2 sensory neuron membrane proteins (SNMPs) (Zeng et al., 2015). Similarly, RNA-Seq analysis of adult antennae identified 90 olfactory-related genes comprising 46 ORs, 15 IRs, 12 OBPs, 15 CSPs, and 2 SNMPs (Liu et al., 2017). Further transcriptomic analysis in 2020 generated 191,974 unigenes with an average length of 791 bp and 289,127 transcripts with an average length of 1,040 bp from, leading to the identification of several genes associated with larval heat acclimation and thermal adaptation (Quan et al., 2020). C. medinalis mitochondrial genome is 15,388 bp long and contains 37 genes arranged similarly to those of other lepidopteran insects (Chai et al., 2012). Earlier transcriptomic assemblies were highly fragmented, limiting comprehensive understanding of the pest’s biology and ecology. However, advances in long-read sequencing technologies, particularly Illumina, PacBio, and Hi-C platforms, enabled chromosome-level genome assembly of C. medinalis. The assembled genome size was approximately 528.5 Mb with 31 chromosomes, 3,248 scaffolds, 4,671 contigs, 38.5% GC content, and 330× chromosomal coverage, leading to the annotation of 15,045 protein-coding genes (Zhao et al., 2021). These genomic advances provide an important foundation for future functional genomics and molecular ecology studies.

 The insect gut plays an important role in nutrient digestion, absorption, immunity, communication, and host defense, while also providing a favorable environment for diverse microbiota, including bacteria, fungi, archaea, protozoa, and viruses  (Dillon and Dillon, 2004; Douglas, 2007; Rajagopal, 2009; Engel and Moran, 2013; Gurung et al., 2019).  Gut microbiota contribute to food digestion, nutrient metabolism, protection against pathogens and predators, and regulation of reproduction and behavior (Engel and Moran, 2013). Subsequently, Illumina MiSeq sequencing identified dominant bacterial phyla such as Actinobacteria, Proteobacteria, Acidobacteria, Firmicutes, and Chloroflexi in fourth-instar larvae (Liu et al., 2016). Further studies across different life stages revealed 22 bacterial phyla, 42 classes, 100 orders, 179 families, 350 genera, and 395 species, with Actinobacteria, Proteobacteria, and Acidobacteria as the predominant groups (Yang et al., 2020). Feeding studies using rice and maize diets demonstrated that host diet significantly influences gut microbial diversity and composition (Yang et al., 2022). Metagenomic analyses further revealed dominance of Firmicutes, particularly Enterococcus spp., including E. gallinarum and E. casseliflavus, accounting for approximately 96% of gut bacteria in different feeding groups (Li et al., 2022).

Management Strategies

Monitoring and forecasting: The identification of the correct C.  medinalis is needed. In adults, the sexual features and patterns of the forewings are required to identify and separate different Cnaphalocrocis species (Islam and Karim, 1997). Conventional visual assessments methods are labor-intensive, time-consuming, and dependent on farmers’ expertise (Martinelli et al., 2015). Consequently, remote sensing (RS) hyperspectral imaging technologies have been increasingly adopted to detect pest-induced changes in crop physiology, canopy structure, pigment composition, and water content over large areas (Huang et al., 2012; Mahlein et al., 2013; Liang et al., 2021). Additional approaches, including soil moisture and crop growth analyses, have also been used to characterize pest habitats (Calderón et al., 2013). Recent studies combined traditional assessment methods with digital tools such as CompuEye and ImageJ program, and leaf symptom analysis software to improve the accuracy of leaf damage estimation caused by C.  medinalis (Adhikari et al., 2023).

C. medinalis monitoring has traditionally relied on searchlight traps, moth-repellent methods, and sex pheromone traps (Wu et al., 2013). However, searchlight traps are often ineffective because C. medinalis adults exhibit weak phototactic responses and urban lights interferes with trap performance traps (Sun et al., 2022; Gao et al., 2023). The disturb-and-count method (DCM) has therefore been widely adopted in Asian countries, although it remains labor-intensive and costly (Yao et al., 2011). The study was also carried out in other countries to confirm that (Z)-13-octadecenyl acetate and (Z)-11-hexadecenyl acetate, while additional compounds have been reported in Japanese populations (Ramachandran et al., 1990). The study was also carried out in other countries to confirm that (Z)-13-octadecenyl acetate and (Z)-11-hexadecenyl acetate, while additional compounds have been reported in Japanese populations (Ramachandran et al., 1990; Wu et al., 2013). Although pheromone traps effectively monitor adult population, they are less reliable for predicting larval density and female fecundity (Zeng et al., 2018). Food attractants such as methyl salicylate, benzaldehyde, valeraldehyde, and phenylacetaldehyde have also shown strong attraction to both male and female moths (Eigenbrode et al., 2016). Integrating food attractants with pheromone-based trapping systems may therefore improve the monitoring and forecasting accuracy of C.  medinalis populations (Zeng et al., 2021).

Cultural control and ecological engineering: Cultural practices are important components of integrated management strategies for C. medinalis. These approaches are environmentally safe cost-effective, and more sustainable than chemical control when implemented on a community-wide scale (Yadav et al., 2012). Recommended practices include crop rotation, destruction of ratoon crops, plowing and flooding after harvest, weed removal, optimum planting time, proper seedbed leveling, balanced plant density, water management, and judicious nitrogen fertilizer application (Devi et al., 2023). Excessive nitrogen fertilization favors C. medinalis outbreaks; therefore, split fertilizer applications at recommended levels are advised (Lella and Jagadeesh, 2023).  Although cultural practices alone may not provide complete control, they are highly effective when integrated with other pest management strategies.

 Ecological engineering has emerged as an important approach for sustainable rice pest management. This concept involves enhancing biodiversity and strengthening natural biological control by regulating agricultural ecosystems (Odum, 1962; Mitsch, 2012). In rice ecosystems, ecological engineering focuses on protecting natural enemies and reducing reliance on pesticides by designing field habitats that favor beneficial arthropods (Gurr et al., 2012). Excessive pesticide application in rice paddies often disrupts predators and parasitic natural enemies, including spiders and parasitic wasps, thereby undermining natural pest control mechanisms (Ngin et al., 2017). Early application of pesticides is particularly harmful because it reduces the effectiveness of late-stage biological control (Ali et al., 2019). Therefore, to protect natural enemies that play a role early in the crop, a "no-spraying-early" strategy is recommended, which avoids pesticide use within the first 30-40 days after sowing or transplanting (Way and Heong, 1994). In addition, maintaining flowering plants along the edges of rice paddies can provide nectar, shelter, and alternative hosts for parasitic wasps and predators, thereby enhancing their effectiveness against C. medinalis and other rice pests (Schoenly et al., 1996).

Biological control: Numerous predators, including hymenopterans, hemipterans, coleopterans, neuropterans, dipterans, spiders, dragonflies, damselflies, coccinellids, carabids, earwigs, and rove beetles, contribute to natural regulation of C.  medinalis populations (Parasappa, 2017). Among them, Spiders, ground beetles, and ladybugs have the highest predation potential, especially during the vegetative growth stage of rice; while mirid bugs and rove beetles are more common during the reproductive growth stage (Bhattacharyya et al., 2006; Rautaray et al., 2019). Dermaptera and Coleoptera insects are particularly effective against the eggs and larvae of C.  medinalis (Meng et al., 2016). In contrast, dragonflies primarily feed on adults and therefore contribute less directly to reducing leaf damage. Many species of predators are useful against C. medinalis throughout the crop-growing period. For instance, predators such as coccinellids, mirid bugs, dragons and damselflies, and carabids could be found throughout the rice crop season. However, coccinellids, spiders, dragonflies, and damselflies were more prominent during the vegetative stage of the rice crop, while Cerambycidae, mirids, and Staphylinids were dominant during the reproductive stage (Rautaray et al., 2019).

 Many species of parasitoids are used as an important biocontrol tool against C. medinalis in China, Pakistan, India, Malaysia, Philippines, Vietnam, Japan, Korea, Sri Lanka, Thailand, Indonesia, Myanmar, Laos, and Nepal (Gurr et al., 2012). There were ninety-one species of hymenopteran parasitoids and fourteen species of dipteran parasitoids have been reported to infect C.  medinalis (Joshi et al., 1987). Since then, many species of parasitoids have been discovered in other countries (Hemachandra and Perera, 2016; Zhu et al., 2022). Significant parasitoid efficiency has been reported in different parts of the world, with most studies focusing on rice-growing regions such as 5–61% in rain-fed regions, 9–13% in dry a regions, and 7–33% in irrigated regions (Litsinger et al., 1987). Among the many parasitic wasps, the efficiency of Trichogramma spp. has been documented in details (Mishra and Kamlesh, 2009; Huang et al., 2012). Among its species, T. chilonis is considered an effective egg parasitoid of C. medinalis after undative releases it ranging from 33 to 89.5 kg/acre of paddy fields.(Khan et al., 2005). The augmentative release of T. chilonis @ 100,000 in the paddy fields of Punjab has been found effective for the significant suppression of C. medinalis (Kaur and Brar, 2008).  Release of T. chilonis @ 100,000/ha, after the application of 1% azadirachtin reduced damage by 41.68 to 98.60% and resulted in an increase of production by 25.79 to 45.13% as compared with the insecticide-treated field (Jena et al., 2012). A field trial revealed the effective dose of T. chilonis among different doses of 50,000, 75,000, 100,000, 125,000, and 150,000 parasitoid eggs /ha, and the results showed that maximum control was achieved at 150,000 parasitized eggs (Zhang et al., 2020). For example, combining the T. chilonis with neem-based products or Bacillus thuringiensis (Bt) can improve the control efficacy against the C. medinalis (Karthikeyan et al., 2007). These combinations are generally more effective because selective biopesticides suppress pest populations while causing relatively little damage to parasitic wasp and predatory natural enemy communities (Jena et al., 2012). In contrast, broad-spectrum insecticides such as monocrotophos and endosulfan, while potentially providing short-term suppression, negatively affect parasitic wasp survival, emergence, and field colonization, thus reducing long-term biocontrol efficiency (Zhang et al., 2020). Despite the promising results achieved by parasitic wasps in controlling the C. medinalis, their large-scale field application remains limited by several practical factors. Large-scale production and storage of high-quality parasitic wasps require specialized facilities and stringent quality control measures to maintain their emergence rate, sex ratio, and field survival rate (Parker et al., 2021). Environmental factors such as temperature, humidity, rainfall, and wind significantly affect the survival rate, dispersal, and field colonization of parasitic wasps after release (Gurr et al., 2012). Furthermore, asynchronous release timing, excessive insecticide applications, and low farmer awareness frequently reduce parasitoid persistence and effectiveness in rice ecosystems (Jena et al., 2012). Consequently, although T. chilonis is an effective component of integrated pest management programs, its long-term success depends on proper release timing, compatibility with selective pesticides, conservation of natural enemies, and coordinated field-level implementation (Karthikeyan et al., 2007; Gurr et al., 2012).

 Entomopathogenic nematodes are important biological control agents against many soil-associated insect pests and are widely distributed worldwide (Kaya and Gaugler, 1993). More than 3,100 natural associations between nematodes and insects involving 23 insect orders and 11 nematode orders have been documented (Kamminga et al., 2012). Several entomopathogenic nematodes species have demonstrated strong pathogenicity against C. medinalis under laboratory conditions such as Steinernema carpocapsae caused up to 98% mortality in fifth-instar larvae of C.  medinalis through rapid multiplication within the host (Srinivas and Prasad, 1993). Similarly, Heterorhabditis indica achieved nearly 100% mortality within 18 to 20 h of exposure (Prasad et al., 2006); while Steinernema asiaticum also showed high virulence against all larval stages under laboratory assays (Padmakumari et al., 2008). Combined inoculation of S. asiaticum and H. indica, increased larval mortality and produced large numbers of infective juveniles, indicating their potential for population persistence and recycling in the field (Sankar et al., 2009). Overall, members of the families Heterorhabditidae and Steinernematidae are considered effective biological control agents against C. medinalis (Devi, 2020). Differences in efficacy among Entomopathogenic nematodes species are mainly associated with their host-seeking behavior, virulence, reproductive capacity, symbiotic bacteria, and tolerance to environmental stress. For example, H. indica is generally more effective because its symbiotic bacteria rapidly kill the host and promote rapid nematode reproduction, while the infectivity and host-finding ability of Steinernemacan vary depending on environmental conditions (Kaya and Gaugler, 1993). Furthermore, the survival rate of infective larvae is strongly influenced by factors such as soil moisture, temperature, ultraviolet radiation, and air humidity, which significantly affect the results of field trials. Despite during laboratory findings, several practical limitations restrict the large-scale application of entomopathogenic nematodes in the control of C. medinalis. Mass production and formulation of infective juveniles require specialized fermentation or in vivo rearing systems, which may increase production costs and reduce commercial feasibility (Shapiro et al., 2012). Their field effectiveness is also highly dependent on proper application timing, particularly during early larval stages and under favorable moisture conditions. Furthermore, exposure to broad-spectrum insecticides and adverse environmental conditions may reduce nematode survival and persistence after application (Naidoo, 2025).

 Entomopathogenic fungi are ubiquitous and have the ability to grow and enter insect bodies after breaching the lining of insect cuticles, thereby contributing to the biological control of both sucking and chewing insect pests in agricultural ecosystems. Although synthetic insecticides are commercially available and widely used, they are not considered permanent solutions because of residue persistence, environmental and health risks, and the development of insecticide resistance (Kalyabina et al., 2021). Consequently, researchers have increasingly focused on biocontrol control agents, including entomopathogenic fungi such as Beauveria bassiana and Metarhizium anisopliae, for the management of C. medinalis (Ullah et al., 2018). In vitro and Greenhouse assay of B. bassiana and M. anisopliae against C. medinalis have demonstrated promising results, with larval mortality reaching up to 70% under laboratory conditions and approximately 50% under greenhouse conditions (Rizwan et al., 2019). A study reported that B. bassiana could serve as an important biological control agent against C. medinalis larvae (Abdullah et al., 2021). Furthermore, the feeding preference of C. medinalis was significantly reduced when plant growth-promoting rhizobacteria were combined with B. bassiana (Karthiba et al., 2010). Studies have also documented the compatibility of entomopathogenic fungi with selected insecticides against C. medinalis (Sahoo and Dangar, 2014). For example, B. bassiana applied at a concentration of 10¹⁰ spores ml^-1 performed effectively when combined with acephate, chlorantraniliprole,  malathion, and thiamethoxam (Sankaranarayanan and Askary, 2017).  In addition, a bioassay revealed that two protein elicitors Hrip 1 and PebB, extracted from Alternaria tenuissima and B. bassiana, respectively, negatively affected the survival, developmental duration, and fecundity of C. medinalis (Basit et al., 2021). Despite these encouraging findings, the efficacy of entomopathogenic fungi varies considerably among fungal species, isolates, and environmental conditions. Beauveria bassiana is often considered more effective than M. anisopliae against foliar-feeding insects such as C. medinalis because it exhibits superior adhesion and penetration abilities on soft-bodied larval cuticles and performs well under humid rice field conditions (Shakeel et al., 2024). In contrast, M. anisopliae exhibit relatively low persistence on plant leaves when exposed to ultraviolet radiation and temperature fluctuation (Alviti et al., 2025). The virulence of entomopathogenic fungi greatly depends on other factors including, enzymatic activity, spore germination rate, tolerant to environmental stress and host specificity (Alviti et al., 2025). Therefore, these factors restrict the field application of entomopathogenic fungi at large-scale in rice fields. Large-scale formulation of fungal spores technically demanding and remain costly, especially in maintaining shelf life and high spore viability during storage and transportation (Mascarin et al., 2024). Entomopathogenic fungi release timing is important, as it is more effective at early stage of C. medinalis larvae, while the older larvae produce thicker cuticles that may exhibit greater toleration and enhanced immune responses (Rizwan et al., 2019). Furthermore, while some insecticides are compatible with entomopathogenic fungi, overuse can inhibit fungal growth, reduce spore germination rates, and negatively affect field persistence (Sahoo and Dangar, 2014).

 Crystal parasporal proteins (Cry) of the gram-positive bacterium B. thuringiensis have been widely used as a foliar spray against agricultural pests worldwide. Cry proteins exhibit insecticidal activity after enzymatic digestion (Heckel, 2020). After ingestion by insects, the Cry protein is activated in the alkaline environment of the midgut, producing toxic fragments. These fragments bind to brush border membrane vesicle receptors, insert into the epithelial cell membrane, leading to ion leakage and cell death (Gurr et al., 2012; Guo et al., 2015). Transgenic rice containing the Bt Cry gene significantly reduced damage caused by lepidopteran pests while also reducing pesticide use (Huang et al., 2005). Studies have shown that Bt rice production can exceed that of non-Bt rice by more than 60% under severe pest pressure (Wang et al., 2010). Furthermore, reduced insecticide use in Bt rice cultivation may improve farmers’ health and decrease environmental contamination (Huang et al., 2015). Several Bt rice lines and Cry proteins, including KMD, Bt Shanyou 63, and Huahui 1 developed through collaborations between Chinese universities and the International Rice Research Institute, have been evaluated for the control of C. medinalis (Chen et al., 2006). Transgenic Bt rice plants with toxicity reduced the suitability for C. medinalis, although resistance and tolerance to Bt toxins have been reported in both laboratory and field populations (Xu et al., 2016; Gassmann et al., 2020; Gothandaraman et al., 2023).  A report suggested that C. medinalis has developed resistance to low concentrations  of Cry proteins, and that Cry1Aa, Cry1Ac, and Cry1C proteins can be degraded by enzymes present in the insect midgut, thereby reducing toxin efficacy (Yang et al., 2023) (Figure 4).  The effectiveness of Cry proteins varies depending on receptor-binding affinity, toxin stability, expression levels in rice tissues, and target insect susceptibility, with Cry1C and Cry2A generally providing broader and stronger toxicity against lepidopteran pests (Bravo et al., 2011). In contrast, some Cry proteins lose efficacy when insects evolve altered midgut receptors or enhanced detoxification mechanisms, leading to reduced toxin activation and increased resistance development (Heckel, 2020). Additionally, the level and tissue-specific expression of Bt toxins in transgenic rice lines strongly influence pest suppression (Chen et al., 2006). Despite the success of Bt rice, several practical constraints limit its large-scale adoption and long-term sustainability. One major challenge is the rapid evolution of insect resistance due to continuous exposure to single Cry toxins, particularly when refuge strategies are not properly implemented (Xu et al., 2016; Gassmann et al., 2020).

Figure 4. The schematic representation shows how Bt toxin affects C.  medinalis larvae.

Host plant resistance: Host plant resistance is an important component of integrated pest management programs for the control of C. medinalis. During the 1990s, extensive efforts were made to evaluate rice germplasm for resistance rice against C. medinalis under both laboratory and field conditions (Heinrichs et al., 1985; Dakshayani et al., 1993). Several resistant rice cultivars and breeding lines were subsequently identified (Khan and Joshi, 1990). By the early 2000s, approximately 271 rice genotypes had been reported to possess resistance against different developmental stages of C. medinalis (Ram et al., 2001). Studies further demonstrated that rice cultivars such as Guanglingxiangjing, Changyou 1, Nanjing 44, Huidao 9, and Huidao 10 exhibited greater resistance than Yangjing 9538, Line 91SP, and TN1 (Xu et al., 2010). Similarly, mutant rice lines MSM 139 and MSM 127 showed significantly higher resistance compared with TN1(Javvaji et al., 2021). Under field conditions, 20 resistant rice germplasm lines were identified among 66 evaluated genotypes, while wild and traditional rice genotypes such as Oryza rhizomatis, TKM 6, Ptb 33, and Oryza minuta also showed resistance by reducing the leaf-folding efficiency of C. medinalis larvae (Punithavalli et al., 2011). In addition, BRRI rice 87 and BRRI rice 72 were identified as resistant cultivars under Kharif-II conditions (Das et al., 2022). Resistance in rice varieties is associated with several biochemical and morphological traits, including higher silica and wax contents, volatile compounds, trichome density, leaf toughness, and plant architecture characteristics such as tiller number and leaf dimensions (Wang et al., 2008). These traits may reduce feeding efficiency, larval establishment, and oviposition preference of C. medinalis, making host plant resistance a cost-effective and environmentally safe strategy, although its effectiveness may vary across environments and pest populations due to genetic variability and adaptation this pest (Punithavalli et al., 2013; Liao et al., 2017). In addition, some resistant cultivars may possess undesirable agronomic traits, such as lower yield potential or poor grain quality, limiting their adoption by farmers. Continuous cultivation of resistant varieties may also impose selection pressure on pest populations, potentially leading to the breakdown of resistance over time (Das et al., 2022).

Chemical control and plant-based insecticides: Heavy damage to rice crops and extensive applications of insecticides against C. medinalis have been widely reported in Pakistan and India (Wakil et al., 2001). Between 1980 to 2000, farmers relied heavily on insecticides such as carbofuron, pyrethroids, chlorpyrifos, malathion, monocrotophos, deltamethrin, diazinon, dimethoate, fenthion, fenitrothion, cypermethrin, endosulfan, methamidophos, carbaryl, and phorate for C. medinalis management (Shaki et al., 2020; Zala and Sipai, 2021). Currently, commonly used insecticides against C. medinalis include abamectin benzoate, cartap, spinosyns, insect growth regulators, fipronil, neonicotinoids, diamides, and neem oil extracts (Shaki et al., 2020). Among them, fipronil and cartap hydrochloride have been extensively used under both laboratory and field conditions because of their strong larvicidal activity (Kumar and Singh, 2013). However, prolonged use of fipronil has reduced its field effectiveness over time, indicating possible resistance development and declining efficacy under outdoor conditions (Supawan and Chongrattanameteekul, 2017). The effectiveness of these insecticides varies depending on application method, dosage, timing, and environmental conditions (Zala and Sipai, 2021). Neonicotinoid insecticides, particularly imidacloprid at 25 g active ingredient per hectare (a. i. ha⁻¹) became widely adopted because of their systemic action and high toxicity against C. medinalis (Krishnaiah et al., 2002). Field reports from southern India indicate an increase in C. medinalis infestations in paddy fields treated with imidacloprid. This suggests that the pesticide-induced pest rebound is related to increased pest reproductive capacity, shortened larval stages, and reduced toxicity to eggs, all of which contribute to rapid population growth (Chintalapati et al., 2016). Similarly, novel diamide insecticides, such as flubendiamide and chlorantraniliprole, initially showed good control efficacy against lepidopteran pests due to their selective action on insect ryanoalkaloid receptor channels (Lahm et al., 2007). Chlorantraniliprole, particularly in combination with thiamethoxam (Virtako), effectively reduced C. medinalis population while conserving predator abundance in paddy ecosystems (Zala and Sipai, 2021). However, the development of resistance to diamide insecticides in C. medinalis populations has raised concerns about their long-term sustainability (Isman, 2020). While chemical insecticides can rapidly and effectively suppress C. medinalis, overuse and repeated use have led to a range of ecological and managerial problems, including insecticide resistance, pest recurrence, environmental pollution, and adverse effects on natural enemies (Chintalapati et al., 2016).

 Plant-derived insecticides are considered ideal alternatives to synthetic insecticides because they are generally safer for non-target organisms and possess multiple mechanisms of action, including repellency, toxicity, antifeedant activity, and growth regulation, making them suitable for controlling agricultural pests (Isman, 2020). Studies have shown that several plant-derived products have significant insecticidal activity against the C. medinalis (Nathan et al., 2005). Several plant-derived insecticides, including commercial formulations and locally available plant extracts, have been evaluated for the control of the C. medinalis (Gopalakrishnan et al., 2014). For example, extracts from Jatropha curcas and Ipomoea carnea have dual antifeedant and larval-killing effects against the rice C. medinalis (Pandey et al., 2017). Similarly, application of Azadirachta indica (neem seed kernel extract) and Vitex negundo (Nirgundi leaf extract), alone, or in combination with Bt, significantly affected the nutritional indicators and feeding behavior of C. medinalis larvae (Gopalakrishnan et al., 2014). Compared with traditional synthetic insecticides, plant-derived insecticides are less likely to cause environmental pollution, pest recurrence, or severe toxicity to beneficial arthropods (Pandey et al., 2017). However, the field efficacy of plant-derived insecticides is generally less stable than that of synthetic chemicals because their activity is affected by plant source, extraction method, environmental conditions, and degradation by sunlight and rainwater (Rajpoot et al., 2021).

Insecticide resistance: Resistance of insects to different synthetic insecticides is an ongoing challenge in pest management worldwide (Sparks et al., 2021). Resistance to insecticides typically involves several detoxifying enzymes that increase detoxification capabilities, to make the insecticide molecules less toxic and more easily eliminated from the body as well as reduce the target site sensitivity (Hilliou et al., 2021). Insecticide resistance in insect pests is a complex process and can be broadly categorized into different types including metabolic resistance, target site resistance, and behavioral resistance (Sparks and Nauen, 2015). Recent research development has been made in identifying resistance genes against different insecticides and their target site mutations affecting their functions (Hawkins et al., 2019). The metabolic resistance occurs through the upregulation of different enzymes such as Cytochrome P450s, Glutathione S-transferases, and Carboxylesterases (Carvalho et al., 2018). Target site resistance of insect-pest occurs due to the mutations in the insecticide target sites that prevent the insecticide molecule from binding, thus ultimately reducing their toxic effects (Renault et al., 2023). Insecticide resistance in C. medinalis was first reported by Rajamma and Das, India in 1969 (Rajamma and Das, 1969).  In 1975, resistance to bromophos-ethyl was documented following repeated field applications (Gangwar, 2015). Subsequent studies in Japan revealed increasing resistance ratios against diazinon, Chlorpyrifos-methyl, dimethylvinophos, tetrachlorvinphos between 1988 and 1991 (Endo, 1993). From  1994 to 2017, C. medinalis populations developed resistance to several insecticide classes under both laboratory and field conditions (Endo et al., 1993; Su et al., 2004; Misra, 2008; Zheng et al., 2011; Zhang et al., 2014; Nayak et al., 2017) (Figure 5).

Figure 5. Timelines on resistance development in C.  medinalis to different insecticides.

In Tamil Nadu, resistance frequencies were particularly high for chlorpyrifos (53.34–80.00%), cartap hydrochloride (51.8–63.34%), and profenophos (43.34–61.34%), while relatively lower resistance levels were observed for chlorantraniliprole (10.66–26.67%) (Shaki et al., 2020) (Figure 6A and B). Moderate resistance levels were also reported for abamectin, spinetoram, and emamectin benzoate (Sun et al., 2023).

Figure 6. Schematic diagram of the C. medinalis Para –orthologous sodium channel representing kdr-associated mutations. Four (I-IV) homologous domains and six (S1-6) transmembrane segments and kdr-associated mutation (K/L) are shown in (A). Schematic representation with red and black dots represents mutation sites identified in C. medinalis, where cylinders show the transmembrane domains are shown in (B).

Genetic and molecular tools: RNA interference (RNAi) has emerged as a promising molecular approach for the management of C. medinalis. In this mechanism, double-stranded RNA (dsRNA) is processed into small interfering RNAs (siRNAs), which bind to complementary mRNA sequences and induce gene silencing (Ma, 2006).  Silencing of certain desired genes may induce insect body deformities, inhibit the chitin metabolism, morbidity, or even cause death (Zhang and Yao, 2017). A study reported that the injection of dsRNA to the larval stage of C.  medinalis decreased the CmHK mRNA level, and that can cause larval and pupal mortality, structural deformities, reduction of average body weight loss, reduced female fecundity, and significant variation in the sex ratio (Shakeel et al., 2020). A study investigated the dsRNA injection knockdown of the target gene (CmCHSB) that is essential for the growth and development of C.  medinalis (Zhang et al., 2021). Knockdown of CmUAP by injection of dsRNA causes stunted growth, reduced feeding and excretion, body weight loss of larvae, and several other developmental disorders (Zhou et al., 2021). A study reveals that the CmdsRNase can degrade the dsRNA and consequently reduce the efficacy of RNAi in C. medinalis; however, co-silencing of CmdsRNase and CmCHS genes can cause adverse effects on the growth and development of C.  medinalis and ultimately improve the RNAi efficiency (Li et al., 2022).

Conclusion: The management of C. medinalis remains challenging due to different factors such as operational constrains, variability, insecticide resistance, and inconsistent field effectiveness. While cultural and biological control, ecological engineering, host plant resistance, and molecular control methods such as RNAi are ecofriendly alternatives, however, their large-scale implantation is limited by issues with long-term stability, implementation difficulty, and with effectiveness. Furthermore, over-reliance on chemical pesticides accelerates resistance development and exacerbates ecological damage. In conclusion, sustainable management of C. medinalis requires an integrated pest control strategy that combines biological, ecological, molecular, and chemical control methods while minimizing environmental risks and resistance development.

Future Research Directions: Future research work on C. medinalis should prioritize climate-adaptive control strategies to reduce reliance on chemical pesticides. The development of predictive models and long term ecological studies are needed to study the pest distribution, outbreaks, and migration under different environmental conditions. The molecular studies such as transcriptomic, genomics, microbiome research, RNAi, should be further utilized to validate different genes associated with heat adaptation, pesticide resistance, reproduction, host selection, and migration. Future biological control such as, predators, parasitoids, entomopathogenic fungi, bacteria, and nematodes research should emphasize field assessments under different agroecological conditions. Ecological engineering approaches, including habitat diversification and pesticide reduction, should also be optimized to protect natural enemies and enhance ecosystem stability. Continuous monitoring of pest adaptation and resistance loss is crucial for maintaining long-term effectiveness. Pesticide resistance management should remain a top priority through research on resistance evolution, cross-resistance, adaptation costs, and molecular markers. Greater emphasis should also be placed on botanical insecticides, pheromone-based systems, remote sensing, hyperspectral imaging, artificial intelligence, and precision agriculture as environmentally safer alternatives to conventional chemical control. Overall, multidisciplinary approaches integrating entomology, molecular biology, ecology, climatology, agronomy, biotechnology, and precision agriculture will be essential for developing sustainable and long-term integrated pest management strategies against C. medinalis.

Acknowledgements: We thank the National Natural Science Foundation of China (Grant Nos. 31360443 and 32060641) for funding this work. We also thank Professor Dr. Khalid Haddi of Federal University of Lavras for his professional English language editing of this manuscript.

Conflict of interest: All authors have read and agreed to the published version of the manuscript. Authors also declare that they have no conflict of interest.

REFERENCES

Abdullah, T., Kuswinanti, T., Aminah, S. N., and Asman, A. (2021). The potential of Beauveria bassiana Vuill. to control rice leaf roller Cnaphalocrocis medinalis Guenée. IOP Conf. Ser.: Earth Environ. Sci. 681(1): 012037. https://doi.org/10.1088/1755-1315/681/1/012037

Adhikari, B., Senapati, R., Mohapatra, M., Mohapatra, L., Nigam, R., and Mohapatra, S. D. (2023). Detection of rice leaf folder, Cnaphalocrocis medinalis (Guenée) (Lepidoptera: Crambidae) infestation using ground-based hyperspectral radiometry. Curr. Sci. 124(8): 964–970. https://doi.org/10.18520/cs/v124/i8/964-970

Ali, M., Bari, M., Haque, S., Kabir, M., Afrin, S., Nowrin, F., and Landis, D. (2019). Establishing next-generation pest control services in rice fields: eco-agriculture. Sci. Rep. 9: 10180. https://doi.org/10.1038/s41598-019-46522-1

Alvi, S. M., Ali, M. A., Chaudhary, S., and Iqbal, S. (2003). Population trends and chemical control of rice leaf folder, Cnaphalocrocis medinalis on rice crop. Int. J. Agric. Biol. 5(4): 615–616.

Alviti Kankanamalage, H. P., Yang, J. Y., Karunarathna, S. C., Tibpromma, S., Kumla, J., Wei, D. P., and Lumyong, S. (2025). Entomopathogenic fungi: insights into recent understanding. J. Microbiol. Biotechnol. 41(6): 179. https://doi.org/10.1007/s11274-025-04377-9

Liao, Q. J., Yang, Y. J., Wang, J., Pang, X., Xu, C. M., Peng, C. L., Lu, Z. X., and Liu, Y. H. (2017). Temperature-dependent development and reproduction of rice leaffolder, Marasmia exigua (Lepidoptera: Pyralidae). PLoS ONE 12(11): e0187972. https://doi.org/10.1371/journal.pone.0187972

Barrion, A. T., Litsinger, J. A., Medina, E. B., Aguda, R. M., Bandong, J. P., Pantua, P. C., Jr., Viajante, V. D., Dela Cruz, C. G., Vega, C. R., Soriano, J. S., Jr., Camanag, E. E., Saxena, R. C., Tryon, E. H., and Shepard, B. M. (1991). The rice Cnaphalocrocis and Marasmia (Lepidoptera: Pyralidae) leaffolder complex in the Philippines: Taxonomy, bionomics and control. Philipp. Entomol. 8(4): 987–1074.

Basit, A., Farhan, M., Essa, M., Abbas, M., Wang, Y., Zhao, D., and Hanan, A. (2021). Effect of two protein elicitors extracted from Alternaria tenuissima and Beauveria bassiana against rice leaf folder (Marasmia exigua). J. King Saud Univ. Sci. 33(8): 101652. https://doi.org/10.1016/j.jksus.2021.101652

Bhattacharyya, B., Basit, A., and Saikia, D. (2006). Parasitoids and predators of rice insect pests of Jorhat district of Assam. J. Biol. Control 20(1): 37–44.

Bodlah, M. A., Gu, L. L., Tan, Y., and Liu, X. D. (2017). Behavioural adaptation of the rice leaf folder Cnaphalocrocis medinalis to short-term heat stress. J. Insect Physiol. 100: 28–34. https://doi.org/10.1016/j.jinsphys.2017.05.004

Bravo, A., Likitvivatanavong, S., Gill, S. S., and Soberón, M. (2011). Bacillus thuringiensis: a story of a successful bioinsecticide. Insect Biochem. Mol. Biol. 41(7): 423–431. https://doi.org/10.1016/j.ibmb.2011.02.006

Calderón, R., Navas-Cortés, J. A., Lucena, C., and Zarco-Tejada, P. J. (2013). High-resolution airborne hyperspectral and thermal imagery for early detection of Verticillium wilt of olive using fluorescence, temperature and narrow-band spectral indices. Remote Sens. Environ. 139: 231–245. https://doi.org/10.1016/j.rse.2013.07.021

Chang, S., Lo, Z., Keng, C., Li, G., Chen, X., and Wu, X. (1980). Studies on the migration of rice leaf roller Cnaphalocrocis medinalis Guenee. Acta Entomol. Sin. 23(2): 130–140.

Carvalho, I. F., Erdmann, L. L., Machado, L. L., Rosa, A., Zotti, M. J., and Neitzke, C. G. (2018). Metabolic resistance in the fall armyworm: an overview. J. Agric. Sci. 10(12): 426–436. https://doi.org/10.5539/jas.v10n12p426

Chai, H.-N., Du, Y. Z., and Zhai, B. P. (2012). Characterization of the complete mitochondrial genomes of Cnaphalocrocis medinalis and Chilo suppressalis (Lepidoptera: Pyralidae). Int. J. Biol. Sci. 8(4): 561. https://doi.org/10.7150/ijbs.4054

Chapman, J. W., Nesbit, R. L., Burgin, L. E., Reynolds, D. R., Smith, A. D., Middleton, D. R., and Hill, J. K. (2010). Flight orientation behaviors promote optimal migration trajectories in high-flying insects. Science 327(5966): 682–685. https://doi.org/10.1126/science.1182999

Chatterjee, M., Chaudhuri, N., Pal, S., and Ghosh, J. (2023). Species composition, occurrence pattern and seasonal incidence of rice leaf folder (Cnaphalocrocis spp.) on kharif rice under Terai agro-ecological conditions of Northern Parts of West Bengal. J. Entomol. Res. 47(2): 386–392.

Chen, M., Zhao, J. Z., Ye, G. Y., Fu, Q., and Shelton, A. M. (2006). Impact of insect-resistant transgenic rice on target insect pests and non-target arthropods in China. Insect Sci. 13(6): 409–420. https://doi.org/10.1111/j.1744-7917.2006.00110.x

Chintalapati, P., Katti, G., Puskur, R. R., and Nagella Venkata, K. (2016). Neonicotinoid-induced resurgence of rice leaffolder, Cnaphalocrocis medinalis (Guénee). Pest Manag. Sci. 72(1): 155–161. https://doi.org/10.1002/ps.4146

Dakshayani, K., Bentur, J., and Kalode, M. (1993). Nature of resistance in rice varieties against leaffolder Cnaphalocrocis medinalis (Guenée). Int. J. Trop. Insect Sci. 14(1): 107–114. https://doi.org/10.1017/S1742758400013473

Dale, D. (1994). Insect pests of the rice plant—their biology and ecology. In: Heinrichs, E. A. (Ed.), Biology and Management of Rice Insects. Wiley Eastern Ltd., New Delhi, India, and International Rice Research Institute (IRRI), Los Baños, Philippines, pp. 363–485.

Das, S., Rahman, M., Sarkar, R., Dash, P. K., Porter, L., and Kamal, M. M. (2022). Characterization of host plant innate immunity from Kharif-II rice cultivars against rice leaf folder (Cnaphalocrocis medinalis Guen.) via in-situ screening and physicochemical assays. J. Asia-Pac. Entomol. 25(1): 101860. https://doi.org/10.1016/j.aspen.2021.101860

Devi, G. (2020). Entomopathogenic nematodes against insect pests of rice. Int. J. Environ. Agric. Biotechnol. 5(4). https://doi.org/10.22161/ijeab.54.34

Devi, D. L., Mandal, K., Dadaro, S., and Kumar, V. (2023). Cultural practices: An urgent need in IPM for sustainable crop production. JA. 78–86. https://doi.org/10.1080/21683565.2012.762438

Dillon, R. J., and Dillon, V. (2004). The gut bacteria of insects: nonpathogenic interactions. Annu. Rev. Entomol. 49(1): 71–92. https://doi.org/10.1146/annurev.ento.49.061802.123416

Douglas, A. E. (2007). Symbiotic microorganisms: untapped resources for insect pest control. Trends Biotechnol. 25(8): 338–342. https://doi.org/10.1016/j.tibtech.2007.06.003

Elanchezhyan, K., and Balakrishnan, N. (2020). Insect pests of rice and their management. In: Meena, M., Kumar, P., Kumar, M., and Meena, R. (Eds.), Recent Trends in Agricultural Sciences, Vol. 1. Akinik Publications, New Delhi, India, pp. 17–35.

Eigenbrode, S. D., Birch, A. N. E., Lindzey, S., Meadow, R., and Snyder, W. E. (2016). A mechanistic framework to improve understanding and applications of push–pull systems in pest management. J. Appl. Ecol. 53(1): 202–212. https://doi.org/10.1111/1365-2664.12592

Endo, S., Tsurumachi, M., and Tanaka, K. (1993). Changes in insecticide susceptibility of the rice leafroller, Cnaphalocrocis medinalis Guenée (Lepidoptera: Pyralidae), collected in Kyushu, and differences in susceptibility between Japanese and Malaysian colonies. Appl. Entomol. Zool. 28(2): 125–130. https://doi.org/10.1303/aez.28.125

Engel, P., and Moran, N. A. (2013). The gut microbiota of insects: diversity in structure and function. FEMS Microbiol. Rev. 37(5): 699–735. https://doi.org/10.1111/1574-6976.12025

Fraenkel, G., Fallil, F., and Kumarasinghe, K. (1981). The feeding behaviour of the rice leaf folder, Cnaphalocrocis medinalis. Entomol. Exp. Appl. 29(2): 147–161.

Fuyan, Z., Chen, X., Xia, Y., Fu, Q., Wang, S., Xu, H., and Zhang, J. (2024). The occurrence characteristics of rice diseases and insect pests and the integration of green control technology in China from 2013 to 2022. Chin. J. Biol. Control. 40(5): 1207.

Fu, X.-W., Li, C., Feng, H. Q., Liu, Z. F., Chapman, J. W., Reynolds, D., and Wu, K. M. (2014). Seasonal migration of Cnaphalocrocis medinalis (Lepidoptera: Crambidae) over the Bohai Sea in northern China. Bull. Entomol. Res. 104(5): 601–609. https://doi.org/10.1017/S0007485314000287

Gangwar, R. K. (2015). Life cycle and abundance of rice leaf folder, Cnaphalocrocis medinalis (Guenee): A review. J. Nat. Sci. Res. 5(15): 103–118.

Gao, L., Yan, R., He, W., and Wu, K. (2023). Use of food attractant to monitor and forecast population dynamics of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae), a long-distance migratory pest. Agronomy 13(8): 2141. https://doi.org/10.3390/agronomy13082141

Gassmann, A. J., Shrestha, R. B., Kropf, A. L., Clair, C. R., and Brenizer, B. D. (2020). Field-evolved resistance by western corn rootworm to Cry34/35Ab1 and other Bacillus thuringiensis traits in transgenic maize. Pest Manag. Sci. 76(1): 268–276. https://doi.org/10.1002/ps.5510

Gothandaraman, R., Sudhakar, D., Varanavasiappan, S., Balakrishnan, N., Udayasuriyan, V., and Natarajan, N. (2023). Adult oviposition preference and larval performance of Cnaphalocrocis medinalis Guenée (Pyralidae: Lepidoptera) on transgenic Bt rice. Int. J. Trop. Insect Sci. 43: 119–129. https://doi.org/10.1007/s42690-023-01026-2

Gopalakrishnan, S., Kumari, B. R., Vijayabharathi, R., Sathya, A., Srinivas, V., and Rao, G. R. (2014). Efficacy of major plant extracts/molecules on field insect pests. In: Advances in Plant Biopesticides, pp. 63–88. https://doi.org/10.1007/978-81-322-1958-3_4

Guo, W., Yang, Y., Xu, J., Xu, H., Zheng, X., and Lv, Z. (2013). Nutritional consumption and utilization by Cnaphalocrocis medinalis larvae of artificial diets with different nitrogen or sugar contents. Chin. J. Appl. Entomol. 50(3): 629–634.

Guo, Z., Kang, S., Chen, D., Wu, Q., Wang, S., Xie, W., and Jurat-Fuentes, J. L. (2015). MAPK signaling pathway alters expression of midgut ALP and ABCC genes and causes resistance to Bacillus thuringiensis Cry1Ac toxin in diamondback moth. PLoS Genet. 11(4): e1005124. https://doi.org/10.1371/journal.pgen.1005124

Guo, J. W., Li, P., Zhang, J., Liu, X. D., Zhai, B. P., and Hu, G. (2019). Cnaphalocrocis medinalis moths decide to migrate when suffering nutrient shortage on the first day after emergence. Insects 10(10): 364. https://doi.org/10.3390/insects10100364

Gurung, K., Wertheim, B., Falcao Salles, J., et al. (2019). The microbiome of pest insects: It is not just bacteria. Entomol. Exp. Appl. 167(3): 156–170. https://doi.org/10.1111/eea.12736

Gurr, G. M., Read, D. M., Catindig, J. L. A., Cheng, J., Liu, J., Lan, L. P., and Heong, K. L. (2012). Parasitoids of the rice leaffolder Cnaphalocrocis medinalis and prospects for enhancing biological control with nectar plants. Agric. For. Entomol. 14(1): 1–12. https://doi.org/10.1111/j.1461-9563.2011.00553.x

Hajjar, M. J., Ahmed, N., Alhudaib, K. A., and Ullah, H. (2023). Integrated insect pest management techniques for rice. Sustainability 15(5): 4499. https://doi.org/10.3390/su15054499

Hawkins, N. J., Bass, C., Dixon, A., and Neve, P. (2019). The evolutionary origins of pesticide resistance. Biol. Rev. 94(1): 135–155. https://doi.org/10.1111/brv.12440

Hilliou, F., Chertemps, T., Maïbèche, M., and Le Goff, G. (2021). Resistance in the genus Spodoptera: Key insect detoxification genes. Insects 12(6): 544. https://doi.org/10.3390/insects12060544

Huang, J., Hu, R., Rozelle, S., and Pray, C. (2005). Insect-resistant GM rice in farmers' fields: Assessing productivity and health effects in China. Science 308(5722): 688–690. https://doi.org/10.1126/science.1108972

Huang, J., Liao, H., Zhu, Y., Sun, J., Sun, Q., and Liu, X. (2012). Hyperspectral detection of rice damaged by rice leaf folder (Cnaphalocrocis medinalis). Comput. Electron. Agric. 82: 100–107. https://doi.org/10.1016/j.compag.2011.12.007

Huang, J., Hu, R., Qiao, F., Yin, Y., Liu, H., and Huang, Z. (2015). Impact of insect-resistant GM rice on pesticide use and farmers’ health in China. Sci. China Life Sci. 58: 466–471. https://doi.org/10.1007/s11427-015-4830-1

Heckel, D. G. (2020). How do toxins from Bacillus thuringiensis kill insects? An evolutionary perspective. Arch. Insect Biochem. Physiol. 104(2): e21673. https://doi.org/10.1002/arch.21673

Heinrichs, E., Camanag, E., and Romena, A. (1985). Evaluation of rice cultivars for resistance to Cnaphalocrocis medinalis Guenee (Lepidoptera: Pyralidae). J. Econ. Entomol. 78(1): 274–278. https://doi.org/10.1093/jee/78.1.274

Hemachandra, K., and Perera, M. (2016). Assessment of the egg parasitoid guild of rice leaf folder, Cnaphalocrocis medinalis (Guenee): A search for biocontrol agents. J. Natl. Sci. Found. Sri Lanka 44(2): 153–160. https://doi.org/10.4038/jnsfsr.v44i2.7999

Heong, K. L. (1993). Rice leaffolders: Are they serious pests? In: Research on Rice Leaffolder Management in China: Proceedings of the International Workshop on Economic Thresholds for Rice Leaffolders in China, 4–6 March 1992, Beijing, China. China National Rice Research Institute, Hangzhou, China, pp. 8–11.

Horgan, F. G. (2017). Insect herbivores of rice: Their natural regulation and ecologically based management. In: Chauhan, B. S., Jabran, K., and Mahajan, G. (Eds.), Rice Production Worldwide. Springer International Publishing, Cham, Switzerland, pp. 279–302. https://doi.org/10.1007/978-3-319-47516-5_12

Islam, Z., and Karim, A. (1997). Leaf folding behaviour of Cnaphalocrocis medinalis (Guenee) and Marasmia patnalis Bradley, and the influence of rice leaf morphology on damage incidence. Crop Prot. 16(3): 215–220. https://doi.org/10.1016/S0261-2194(96)00082-3

Isman, M. B. (2020). Botanical insecticides in the twenty-first century—fulfilling their promise? Annu. Rev. Entomol. 65: 233–249. https://doi.org/10.1146/annurev-ento-011019-025010

Javvaji, S., Telugu, U. M., Venkata, R. D. B., Madhav, M. S., Rathod, S., and Chintalapati, P. (2021). Characterization of resistance to rice leaf folder, Cnaphalocrocis medinalis, in mutant Samba Mahsuri rice lines. Entomol. Exp. Appl. 169(9): 859–875. https://doi.org/10.1111/eea.13071

Jena, B., Gupta, S., and Das, S. (2012). Effectiveness of integrated pest management modules in suppression of major insect pests in rice. Plant Prot. Res. J. 9(1): 57–59.

Joshi, R. C., Cadapan, E. A., and Heinrichs, E. A. (1987). Natural enemies of rice leaf folder Cnaphalocrocis medinalis (Guenée)—a critical review. IRRI Res. Pap. Ser. 126: 1–31.

Junaid, K., and Khan, S. A. (2023). Compatible and non-compatible interaction of rice germplasms against rice leaf folder Cnaphalocrocis medinalis (Guenee) (Lepidoptera: Pyralidae). Pak. J. Zool. 55(6): 2727–2735. https://doi.org/10.17582/journal.pjz/20211004091059

Kamminga, K. L., Davis, J. A., Stock, S. P., and Richter, A. R. (2012). First report of a mermithid nematode infecting Piezodorus guildinii and Acrosternum hilare (Hemiptera: Pentatomidae) in the United States. Florida Entomol. 95(1): 214–217. https://doi.org/10.1653/024.095.0137

Kalyabina, V. P., Esimbekova, E. N., Kopylova, K. V., and Kratasyuk, V. A. (2021). Pesticides: formulants, distribution pathways and effects on human health. Toxicol. Rep. 8: 1179–1192. https://doi.org/10.1016/j.toxrep.2021.06.004

Karthiba, L., Saveetha, K., Suresh, S., Raguchander, T., Saravanakumar, D., and Samiyappan, R. (2010). PGPR and entomopathogenic fungus bioformulation for the synchronous management of leaffolder pest and sheath blight disease of rice. Pest Manag. Sci. 66(5): 555–564. https://doi.org/10.1002/ps.1907

Kaur, R., and Brar, K. S. (2008). Evaluation of different doses of Trichogramma species for the management of leaf folder and stem borer on basmati rice. J. Biol. Control 22(1): 131–135. https://doi.org/10.18311/jbc/2008/3809

Kaya, H. K., and Gaugler, R. (1993). Entomopathogenic nematodes. Annu. Rev. Entomol. 38(1): 181–206. https://doi.org/10.1146/annurev.en.38.010193.001145

Khan, Z. R., and Joshi, R. C. (1990). Varietal resistance to Cnaphalocrocis medinalis (Guenee) in rice. Crop Prot. 9(4): 243–251. https://doi.org/10.1016/0261-2194(90)90056-J

Khan, R. A., Khan, J. A., Jamil, F., and Hamed, M. (2005). Resistance of different basmati rice varieties to stem borers under different control tactics of IPM and evaluation of yield. Pak. J. Bot. 37(2): 319–324.

Karthikeyan, K., Jacob, S., and Purushothaman, S. (2007). Field evaluation of egg parasitoids, Trichogramma japonicum Ashmead and Trichogramma chilonis Ishii, against rice yellow stem borer and leaf folder. J. Biol. Control. 21(2): 261–265.

Ki-Baik, U. (2006). Rearing system for rice leaffolder, Cnaphalocrocis medinalis (Lepidoptera: Crambidae) using corn seedlings. Korean J. Appl. Entomol. 45(1): 91–95.

Krishnaiah, N., Pasalu, I. C., Ramaprasad, A., Lingaiah, T., Lakshminarayanamma, V., Raju, G., and Verma, N. R. (2002). Relative efficacy of newer insecticides against insect pests of rice under field conditions. Pestology 26: 17–20.

Kumar, R., and Singh, B. (2013). Persistence and metabolism of fipronil in rice (Oryza sativa Linnaeus) field. Bull. Environ. Contam. Toxicol. 90: 482–488. https://doi.org/10.1007/s00128-013-0955-0

Lahm, G. P., Stevenson, T. M., Selby, T. P., Freudenberger, J. H., Dubas, C. M., Smith, B. K., and Bellin, C. A. (2007). Rynaxypyr™: A new anthranilic diamide insecticide acting at the ryanodine receptor. In: Ohkawa, H., Miyagawa, H., and Lee, P. W. (Eds.), Pesticide Chemistry: Crop Protection, Public Health, Environmental Safety. Wiley-VCH, Weinheim, Germany, pp. 111–120. https://doi.org/10.1002/9783527611249.ch11

Lella, N., and Jagadeesh, K. (2023). Ecological IPM strategies for management of pests of rice. Pharma Innov. J. 12(5): 3814–3818. https://doi.org/10.56424/pijp.2023.12.5.466

Lee, S. C. (1992). Towards integrated pest management of rice in Korea. Korean J. Appl. Entomol. 31(3): 205–240.

Li, C., Han, G., Sun, J., Huang, L., Lu, Y., Xia, Y., and Xu, J. (2022). The gut microbiota composition of Cnaphalocrocis medinalis and their predicted contribution to larval nutrition. Front. Microbiol. 13: 909863. https://doi.org/10.3389/fmicb.2022.909863

Li, J., Du, J., Li, S., and Wang, X. (2022). Identification and characterization of a double-stranded RNA degrading nuclease influencing RNAi efficiency in the rice leaf folder Cnaphalocrocis medinalis. Int. J. Mol. Sci. 23(7): 3961. https://doi.org/10.3390/ijms23073961

Li, S. W., Yang, H., Liu, Y. F., Liao, Q. R., Du, J., and Jin, D. C. (2012). Transcriptome and gene expression analysis of the rice leaf folder, Cnaphalocrocis medinalis. PLoS ONE 7(11): e47401. https://doi.org/10.1371/journal.pone.0047401

Liang, G. C., Ouyang, Y. C., and Dai, S. M. (2021). Detection and classification of rice infestation with rice leaf folder (Cnaphalocrocis medinalis) using hyperspectral imaging techniques. Remote Sens. 13(22): 4587. https://doi.org/10.3390/rs13224587

Liao, C. T., and Chen, C. L. (2017). Oviposition preference and larval performance of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) on rice genotypes. J. Econ. Entomol. 110(3): 1291–1297. https://doi.org/10.1093/jee/tox095

Liu, X., Yang, Y., Liao, Q., Xu, H., Liu, Y., and Lu, Z. (2016). Analysis of the bacterial community structure and diversity in the intestine of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). Acta Entomol. Sin. 59(9): 965–976.

Liu, S., Wang, W. L., Zhang, Y. X., Zhang, B. X., Rao, X. J., Liu, X. M., and Li, S. G. (2017). Transcriptome sequencing reveals abundant olfactory genes in the antennae of the rice leaffolder, Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). Entomol. Sci. 20(1): 177–188. https://doi.org/10.1111/ens.12230

Liu, H., Han, Y., Zhong, Y., Hou, A., and Ali, M. (2021). Preference and performance of the rice leaf folder, Cnaphalocrocis medinalis, in relation to rice developmental stage. Entomol. Exp. Appl. 169(8): 701–710. https://doi.org/10.1111/eea.13051

Litsinger, J., Barrion, A. T., and Soekarna, D. (1987). Upland Rice Insect Pests: Their Ecology, Importance and Control. International Rice Research Institute (IRRI), Los Baños, Philippines.

Litsinger, J. A. (1994). Cultural, mechanical and physical control of rice insects. In: Heinrichs, E. A. (Ed.), Biology and Management of Rice Insects. International Rice Research Institute (IRRI), Los Baños, Philippines, pp. 549–584.

Litsinger, J. A., Bandong, J. B., Canapi, B. L., Dela Cruz, C. G., Pantua, P. C., Alviola, A. L., and Batay, I. E. H. (2005). Evaluation of action thresholds for chronic rice insect pests in the Philippines. I. Less frequently occurring pests and overall assessment. Int. J. Pest Manag. 51(1): 45–61. https://doi.org/10.1080/09670870500070936

Lv, W., Jiang, X., Chen, X., Cheng, Y., Xia, J., and Zhang, L. (2021). Flight and reproduction variations of rice leaf roller, Cnaphalocrocis medinalis, in response to different rearing temperatures. Insects 12(12): 1083. https://doi.org/10.3390/insects12121083

Ma, V. S. (2006). Control of translation and mRNA degradation by miRNAs and siRNAs. Genes Dev. 20(5): 515–524. https://doi.org/10.1101/gad.1399806

Maclean, J. L., Hardy, B., and Hettel, G. P. (2013). Rice Almanac: Source Book for One of the Most Important Economic Activities on Earth. 4th Ed. International Rice Research Institute (IRRI), Los Baños, Philippines.

Mahlein, A. K., Rumpf, T., Welke, P., Dehne, H. W., Plümer, L., Steiner, U., and Oerke, E. C. (2013). Development of spectral indices for detecting and identifying plant diseases. Remote Sens. Environ. 128: 21–30. https://doi.org/10.1016/j.rse.2012.10.019

Martinelli, F., Scalenghe, R., Davino, S., Panno, S., Scuderi, G., Ruisi, P., Villa, P., Stroppiana, D., Boschetti, M., Goulart, L. R., and Davis, C. (2015). Advanced methods of plant disease detection: A review. Agron. Sustain. Dev. 35: 1–25. https://doi.org/10.1007/s13593-014-0246-1

Mashhoor, K., Ramesh, N., Azar, K., and Shanas, S. (2019). Molecular phylogeny of rice leaf folder (Cnaphalocrocis medinalis). Res. J. Biotechnol. 14(8): 1–8.

Mascarin, G. M., Golo, P. S., de Souza Ribeiro-Silva, C., Muniz, E. R., de Oliveira Franco, A., Kobori, N. N., and Fernandes, É. K. K. (2024). Advances in submerged liquid fermentation and formulation of entomopathogenic fungi. Appl. Microbiol. Biotechnol. 108(1): 451. https://doi.org/10.1007/s00253-023-12981-4

Meng, J., Mabubu, J. I., Han, Y., He, Y., Zhao, J., Hua, H., and Wu, G. (2016). No impact of transgenic Cry1C rice on the rove beetle Paederus fuscipes, a generalist predator of brown planthopper Nilaparvata lugens. Sci. Rep. 6: 30303. https://doi.org/10.1038/srep30303

Mitsch, W. J. (2012). What is ecological engineering? Ecol. Eng. 45: 5–12. https://doi.org/10.1016/j.ecoleng.2012.04.013

Mishra, D., and Kamlesh, K. (2009). Field efficacy of bio-agent Trichogramma spp. against stem borer and leaf folder in rice crop under mid-western plain zone of Uttar Pradesh. Environ. Ecol. 27(4A): 1885–1887.

Misra, H. P. (2008). Management of the rice leaffolder, Cnaphalocrocis medinalis (Guenee) by newer insecticides. Oryza 45(3): 252–254.

Morshed, M. N., Uddin, M. E. M. H. R., and Sultana, N. (2020). Effect of temperature, rainfall and relative humidity on seasonal incidence of major rice insect pests. Int. J. Biosci. 17: 92–102.

Morshed, M. N., Mamun, M. A. A., Nihad, S. A. I., Rahman, M. M., Sultana, N., and Rahman, M. M. (2023). Effect of weather variables on seasonal abundance of rice insects in southeast coastal region of Bangladesh. J. Agric. Food Res. 11: 100513. https://doi.org/10.1016/j.jafr.2023.100513

Naidoo, K. (2025). Agricultural pest management: Identification, characterisation, and stress tolerance of entomopathogenic nematodes. M.Sc. Thesis, University of the Witwatersrand, Johannesburg, South Africa.

Nathan, S. S., Chung, P. G., and Murugan, K. (2005). Effect of biopesticides applied separately or together on nutritional indices of the rice leaffolder Cnaphalocrocis medinalis. Phytoparasitica 33: 187–195. https://doi.org/10.1007/BF02979825

Nayak, S. B., Manoharan, T., and Soundararajan, R. (2017). Monitoring insecticide resistance to leaffolder, Cnaphalocrocis medinalis (Guenee) (Lepidoptera: Pyralidae) in rice growing regions of Tamil Nadu. J. Entomol. Zool. Stud. 5: 23–26.

Ngin, C., Suon, S., Tanaka, T., Yamauchi, A., Kawakita, K., and Chiba, S. (2017). Impact of insecticide applications on arthropod predators and plant feeders in Cambodian rice fields. Phytobiomes J. 1(3): 128–137. https://doi.org/10.1094/PBIOMES-05-17-0017-R

Odum, H. T. (1962). Man in the ecosystem. In: Proceedings of the Lockwood Conference on the Suburban Forest and Ecology. Bull. Conn. Agric. Exp. Stn. 652: 57–75.

Padmavathi, C., Katti, G., Padmakumari, A., Voleti, S., and Subba Rao, L. (2013). The effect of leaffolder Cnaphalocrocis medinalis (Guenee) [Lepidoptera: Pyralidae] injury on plant physiology and yield loss in rice. J. Appl. Entomol. 137(4): 249–256. https://doi.org/10.1111/j.1439-0418.2011.01688.x

Parasappa, H. (2017). Rice insect pests and their natural enemies complex in different rice ecosystems of Cauvery command areas of Karnataka. J. Entomol. Zool. Stud. 5(5): 335–338.

Parker, A. G., Mamai, W., and Maiga, H. (2021). Mass-rearing for the sterile insect technique. In: Dyck, V. A., Hendrichs, J., and Robinson, A. S. (Eds.), Sterile Insect Technique. CRC Press, Boca Raton, FL, USA, pp. 283–316. https://doi.org/10.1201/9781003035572-9

Punithavalli, M., Muthukrishnan, N., and Balaji, R. (2011). Influence of rice leaf morphology on the folding characteristics of rice leaffolder, Cnaphalocrocis medinalis. Indian J. Plant Prot. 39(2): 93–99.

Punithavalli, M., Muthukrishnan, N., and Rajkumar, M. B. (2013). Influence of rice genotypes on folding and spinning behaviour of leaffolder (Cnaphalocrocis medinalis) and its interaction with leaf damage. Rice Sci. 20(6): 442–450. https://doi.org/10.1016/S1672-6308(13)60158-7

Padmakumari, A., Katti, G., Sankar, M., and Prasad, J. (2008). Entomopathogenic Nematodes in Rice Pest Management. Technical Bulletin No. 32. Directorate of Rice Research, Hyderabad, India.

Pandey, N., Dubey, V., and Gajbhiye, R. K. (2017). Comparative potential of different plant products and synthetic insecticides and their economics against Cnaphalocrocis medinalis in rice. Int. J. Curr. Microbiol. Appl. Sci. 6(5): 2501–2508. https://doi.org/10.20546/ijcmas.2017.605.280

Pathak, M. D., and Khan, Z. R. (1994). Insect Pests of Rice. International Rice Research Institute (IRRI), Los Baños, Philippines.

Prasad, J., Katti, G., Padmakumari, A., and Sankar, M. (2006). Bioefficacy of indigenous entomopathogenic nematodes against leaf folder, Cnaphalocrocis medinalis Guenee in rice. In: Proceedings of the 2nd International Congress of Nematology, 9–13 October 2006, pp. 1–5.

Quan, P. Q., Li, M. Z., Wang, G. R., Gu, L. L., and Liu, X. D. (2020). Comparative transcriptome analysis of the rice leaf folder, Cnaphalocrocis medinalis, to heat acclimation. BMC Genomics 21(1): 1–12. https://doi.org/10.1186/s12864-020-6741-8

Rajamma, P., and Das, N. (1969). Studies on the biology and control of the rice leaf roller Cnaphalocrocis medinalis Guen. 110–112.

Rajagopal, R. (2009). Beneficial interactions between insects and gut bacteria. Indian J. Microbiol. 49: 114–119. https://doi.org/10.1007/s12088-009-0023-z

Rajpoot, S. K. S., Dixit, S., Prasad, V., Giri, S. P., Singh, R. A., Parkash, N., Upadhay, A. L., and Chandra, S. (2021). Evaluation of insecticides and biopesticides against Scirpophaga incertulas (Walker), leaf folder (Cnaphalocrocis medinalis) and green leafhopper in basmati rice. Pharma Innov. J. 10(12, Spl. Issue): 368–371.

Ramachandran, R., Caballero, P., and Khan, Z. (1990). Pheromone components of rice leaffolders (Cnaphalocrocis medinalis and Marasmia patnalis). J. Entomol. Res. 15(4): 25–26.

Ram, L., Singh, R., and Singh, R. (2001). Sources and mechanisms of resistance in rice against rice leaffolder Cnaphalocrocis medinalis (Guenee). Agric. Rev. 22(1): 1–12.

Ranasinghe, M. A. S. K. (1992). Paddy pests in Sri Lanka. Sci. Educ. Ser. 30: 47–59.

Rani, W. B., Amutha, R., Muthulakshmi, S., Indira, K., and Mareeswari, P. (2007). Diversity of rice leaf folders and their natural enemies. Res. J. Agric. Biol. Sci. 3(5): 394–397.

Rautaray, B., Bhattacharya, S., Panigrahi, D., and Dash, S. (2019). Studies on species diversity of rice leaf folder and their natural enemies in north eastern coastal plains of Odisha. Int. J. Curr. Microbiol. Appl. Sci. 8(10): 634–645. https://doi.org/10.20546/ijcmas.2019.810.071

Renault, D., Elfiky, A., and Mohamed, A. (2023). Predicting the insecticide-driven mutations in a crop pest insect: Evidence for multiple polymorphisms of acetylcholinesterase gene with potential relevance for resistance to chemicals. Sci. Rep. 13: 18937–18955. https://doi.org/10.1038/s41598-023-45809-2

Rizwan, M., Atta, B., Sabir, A. M., Yaqub, M., and Qadir, A. (2019). Evaluation of the entomopathogenic fungi as a non-traditional control of the rice leaf roller, Cnaphalocrocis medinalis (Guenee) (Lepidoptera: Pyralidae) under controlled conditions. Egypt. J. Biol. Pest Control 29(1): 1–4. https://doi.org/10.1186/s41938-019-0121-2

Roseli, M., Adam, N. A., Hong, L. W., and Yaakop, S. (2021). Population fluctuation of rice leaffolder, Cnaphalocrocis medinalis, in two consecutive rice seasons. J. Agrobiotechnol. 12(1): 10–22. https://doi.org/10.37231/jab.2021.12.1.245

Sahoo, B., and Dangar, T. K. (2014). Compatibility of some fungal entomopathogens of rice leaf folder (Cnaphalocrocis medinalis Guenee) with selected chemical insecticides. J. Microbiol. Biotechnol. Res. 4(4): 1–7. https://doi.org/10.13140/RG.2.2.31748.83840

Sankaranarayanan, C., and Askary, T. H. (2017). Status of entomopathogenic nematodes in integrated pest management strategies in India. In: Askary, T. H., and Martinelli, P. R. P. (Eds.), Biocontrol Agents: Entomopathogenic and Slug Parasitic Nematodes, pp. 362–382.

Sankar, M., Prasad, J., Padmakumari, A., Katti, G., and Divya, K. (2009). Combined application of two entomopathogenic nematodes, Heterorhabditis indica and Steinernema asiaticum to control the rice leaf folder, Cnaphalocrocis medinalis (Guenee). J. Biopest. 2(2): 135–140.

Schoenly, K. G., Cohen, J. E., Heong, K., Arida, G. S., Barrion, A. T., and Litsinger, J. A. (1996). Quantifying the impact of insecticides on food web structure of rice-arthropod populations in a Philippine farmer’s irrigated field: A case study. In: Polis, G. A., and Winemiller, K. O. (Eds.), Food Webs: Integration of Patterns and Dynamics. Springer, New York, USA, pp. 343–351. https://doi.org/10.1007/978-1-4612-3985-2_33

Shakeel, M., Du, J., Li, S. W., Zhou, Y. J., Sarwar, N., and Bukhari, S. A. H. (2020). Characterization, knockdown and parental effect of hexokinase gene of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) revealed by RNA interference. Genes 11(11): 1258. https://doi.org/10.3390/genes11111258

Shakeel, M., Khan, A., Du, J., Basit, A., Yang, G., Haddi, K., and Li, S. (2024). Knockdown of the glucosamine-6-phosphate N-acetyltransferase gene by RNA interference enhances the virulence of entomopathogenic fungi against rice leaffolder Cnaphalocrocis medinalis. Pestic. Biochem. Physiol. 205: 106119. https://doi.org/10.1016/j.pestbp.2024.106119

Shaki, S., Morshed, M. N., Sultana, N., and Das, G. (2020). Comparative efficacy of some newer insecticides against rice leaf folder (Cnaphalocrocis medinalis Guen.) and their effects on natural enemies in rice ecosystem. Trends Agric. Res. Biotechnol. 1(2): 77–81.

Shapiro-Ilan, D. I., Gaugler, R., and Lewis, W. (2012). Mass production and in vivo production of entomopathogenic nematodes. J. Nematol. 44(2): 206–217. https://doi.org/10.21307/jofnem-2017-015

Shono, Y., and Hirano, M. (1989). Improved mass-rearing of the rice leaffolder, Cnaphalocrocis medinalis (Guenee) (Lepidoptera: Pyralidae) using corn seedlings. Appl. Entomol. Zool. 24(3): 258–263. https://doi.org/10.1303/aez.24.258

Sparks, T. C., and Nauen, R. (2015). IRAC: Mode of action classification and insecticide resistance management. Pestic. Biochem. Physiol. 121: 122–128. https://doi.org/10.1016/j.pestbp.2014.11.014

Sparks, T. C., Storer, N., Porter, A., Slater, R., and Nauen, R. (2021). Insecticide resistance management and industry: The origins and evolution of the Insecticide Resistance Action Committee (IRAC) and the mode of action classification scheme. Pest Manag. Sci. 77(6): 2609–2619. https://doi.org/10.1002/ps.6254

Srinivas, P., and Prasad, J. S. (1993). Record of DD-136 nematode infection on rice leaf-folder (Cnaphalocrocis medinalis). J. Biol. Control 7(2): 139–140.

Su, J. K., Chu, B., and Chen, W. M. (2003). Preliminary study on method of determining rice leaf roller resistance to insecticides and resistance monitoring. Acta Agric. Shanghai 19(4): 81–84.

Sun, G., Liu, S., Luo, H., Feng, Z., Yang, B., Luo, J., and Xu, J. (2022). Intelligent monitoring system of migratory pests based on searchlight trap and machine vision. Front. Plant Sci. 13: 897739. https://doi.org/10.3389/fpls.2022.897739

Sun, Y., Liu, S. T., Ling, Y., Wang, L., Ni, H., Guo, D., and Gao, C. F. (2023). Insecticide resistance monitoring of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) and its mechanism to chlorantraniliprole. Pest Manag. Sci. 79(9): 3290–3299. https://doi.org/10.1002/ps.7556

Supawan, J., and Chongrattanameteekul, W. (2017). Influence of humidity, rainfall and fipronil toxicity on rice leaffolder (Cnaphalocrocis medinalis). Sci. Asia 43: 82–87. https://doi.org/10.2306/scienceasia1513

Tiwari, S., Chaudhary, J., Puranik, H., and Singh, B. (2021). Influence of weather parameters on population dynamics of rice leaf folder (RLF) in rice crop at Raipur district of Chhattisgarh. J. Entomol. Zool. Stud. 9: 1242–1248.

Ullah, M. I., Arshad, M., Abdullah, A., Khalid, S., Iftikhar, Y., and Zahid, S. M. A. (2018). Use of the entomopathogenic fungi Beauveria bassiana (Hyphomycetes: Moniliales) and Isaria fumosorosea (Hypocreales: Cordycipitaceae) to control Diaphorina citri Kuwayama (Hemiptera: Liviidae) under laboratory and semi-field conditions. Egypt. J. Biol. Pest Control 28: 1–5. https://doi.org/10.1186/s41938-018-0047-0

Wakil, W., Akbar, R., and Gulzar, A. (2001). Evaluation of different insecticides against rice stem borer and rice leaf folder. Pak. Entomol. 23(1–2): 49–50.

Waldbauer, G. P., and Marciano, A. P. (1981). Mass rearing of the rice leaf folder, Cnaphalocrocis medinalis Guenee (Lepidoptera: Pyralidae) under greenhouse conditions. Int. Rice Res. Newsl. 6(3): 1–8.

Walker, F. (1859). List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, Part XVII. British Museum, London, UK, pp. 255–508.

Wang, Q., Xu, L., and Wu, J. C. (2008). Physical and biochemical mechanisms of resistance of different rice varieties to the rice leaf folder, Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). Acta Entomol. Sin. 51(12): 1265–1270.

Wang, F., Zhang, X., and Zhai, B. P. (2010). Flight and re-migration capacity of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée) (Lepidoptera: Crambidae). Acta Entomol. Sin. 53(11): 1265–1272.

Wang, Y., Zhang, S., Ren, X., and Su, J. (2013). Optimization of an artificial diet for the rice leaf-folder, Cnaphalocrocis medinalis. Chin. J. Appl. Entomol. 50(3): 635–640.

Wang, Y. C., Zhang, S. K., Ren, X. B., and Su, J. (2014). Effects of dietary additives in artificial diets on survival and larval development of Cnaphalocrocis medinalis (Lepidoptera: Crambidae). Fla. Entomol. 97(3): 1041–1048. https://doi.org/10.1653/024.097.0312

Wang, F. Y., Yang, F., Lu, M. H., Luo, S. Y., Zhai, B. P., Lim, K. S., and Hu, G. (2017). Determining the migration duration of rice leaf folder (Cnaphalocrocis medinalis (Guenée)) moths using a trajectory analytical approach. Sci. Rep. 7(1): 39853. https://doi.org/10.1038/srep39853

Way, M. J., and Heong, K. L. (1994). The role of biodiversity in the dynamics and management of insect pests of tropical irrigated rice: A review. Bull. Entomol. Res. 84(4): 567–587. https://doi.org/10.1017/S0007485300032871

Wu, J., Wu, X., Chen, H., Xu, L., Liu, G., Mao, B., and Du, Y. (2013). Optimization of the sex pheromone of the rice leaffolder moth Cnaphalocrocis medinalis as a monitoring tool in China. J. Appl. Entomol. 137(7): 509–518. https://doi.org/10.1111/jen.12018

Xu, J., Wang, Q. X., and Wu, J. C. (2010). Resistance of cultivated rice varieties to Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). J. Econ. Entomol. 103(4): 1166–1171. https://doi.org/10.1603/EC09331

Xu, Y., Li, X., Chen, F., Zhai, B. P., Hou, M. L., and Han, L. Z. (2013). Analysis of nutritional components of rice leaves and its application in the development of an artificial diet for Cnaphalocrocis medinalis. Chin. J. Appl. Entomol. 50(3): 641–650.

Xu, L., Pan, Z. Z., Zhang, J., Liu, B., Zhu, Y. J., and Chen, Q. X. (2016). Proteolytic activation of Bacillus thuringiensis Cry2Ab through a belt-and-braces approach. J. Agric. Food Chem. 64(38): 7195–7200. https://doi.org/10.1021/acs.jafc.6b02513

Yadav, M. K., Yadav, A., Singh, A. K., Mahajan, G., Singh, M. K., Singh, R., and Babu, S. (2012). Ridge planted pigeonpea and furrow planted rice in an intercropping system as affected by nitrogen and weed management. In: Crop Production Technologies. IntechOpen.

Yadav, S., and Kumar, V. (2018). Feeding the world while caring for the planet. Direct Seed Rice Consort. Newsl. 1(2): 3–4.

Yang, Y., Liu, X., Xu, H., Liu, Y., Ali, P., Bodlah, M. A., and Lu, Z. (2020). The abundance and diversity of gut bacteria of rice leaffolder Cnaphalocrocis medinalis (Guenée) across life stages. J. Asia-Pac. Entomol. 23(2): 430–438. https://doi.org/10.1016/j.aspen.2020.01.008

Yang, Y., Liu, X., Xu, H., Liu, Y., and Lu, Z. (2022). Effects of host plant and insect generation on shaping of the gut microbiota in the rice leaffolder, Cnaphalocrocis medinalis. Front. Microbiol. 13: 824224. https://doi.org/10.3389/fmicb.2022.824224

Yang, Y., Wu, Z., He, X., Xu, H., and Lu, Z. (2023). Processing properties and potency of Bacillus thuringiensis Cry toxins in the rice leaffolder Cnaphalocrocis medinalis (Guenée). Toxins 15(4): 275. https://doi.org/10.3390/toxins15040275

Yao, S., Wu, J., Zheng, Y., Jin, Z., Lu, Z., and Hu, J. (2011). Application of sex pheromone of the rice leaf folder, Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) in monitoring its population. Acta Entomol. Sin. 54(4): 490–494.

Yu, H. Z., Wen, D. F., Wang, W. L., Geng, L., Zhang, Y., and Xu, J. P. (2015). Identification of genes putatively involved in chitin metabolism and insecticide detoxification in the rice leaf folder (Cnaphalocrocis medinalis) larvae through transcriptomic analysis. Int. J. Mol. Sci. 16(9): 21873–21896. https://doi.org/10.3390/ijms160921873

Zala, M., and Sipai, S. (2021). Efficacy of insecticides against rice leaf folder, Cnaphalocrocis medinalis (Guenee). Pharma Innov. 10(12): 874–879.

Zeng, F. F., Zhao, Z. F., Yan, M. J., Zhou, W., Zhang, Z., Zhang, A., and Wang, M. Q. (2015). Identification and comparative expression profiles of chemoreception genes revealed from major chemoreception organs of the rice leaf folder, Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). PLoS One 10(12): e0144267. https://doi.org/10.1371/journal.pone.0144267

Zeng, F. F., Liu, H., Zhang, A., Lu, Z. X., Leal, W. S., Abdelnabby, H., and Wang, M. Q. (2018). Three chemosensory proteins from the rice leaf folder Cnaphalocrocis medinalis involved in host volatile and sex pheromone reception. Insect Mol. Biol. 27(6): 710–723. https://doi.org/10.1111/imb.12511

Zeng, J., Zhang, T., Wang, L., Wu, Q., Lu, M., and Wu, K. (2021). Preliminary application of food attractant trapping in monitoring population dynamics of rice leaf folder, Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) in China. Plant Prot. 47: 203–214.

Zhao, X., Xu, H., He, K., Shi, Z., Chen, X., Ye, X., and Gao, L. (2021). A chromosome-level genome assembly of rice leaffolder, Cnaphalocrocis medinalis. Mol. Ecol. Resour. 21(2): 561–572. https://doi.org/10.1111/1755-0998.13278

Zhang, S. K., Ren, X. B., Wang, Y. C., and Su, J. (2014). Resistance in Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) to new chemistry insecticides. J. Econ. Entomol. 107(2): 815–820. https://doi.org/10.1603/EC13447

Zhang, L., Pan, P., Sappington, T. W., Lu, W., Luo, L., and Jiang, X. (2015). Accelerated and synchronized oviposition induced by flight of young females may intensify larval outbreaks of the rice leaf roller. PLoS One 10(3): e0121821. https://doi.org/10.1371/journal.pone.0121821

Zhang, L., Lu, H., Guo, K., Yao, S., and Cui, F. (2017). Insecticide resistance status and detoxification enzymes of wheat aphids Sitobion avenae and Rhopalosiphum padi. Sci. China Life Sci. 60(8): 927–930. https://doi.org/10.1007/s11427-017-9095-5

Zhang, X., Li, Y., Wang, H., Chen, J., and Zhou, L. (2020). Field evaluation of Trichogramma chilonis release density for management of lepidopteran pests in rice ecosystems. Biol. Control 150: 104362. https://doi.org/10.1016/j.biocontrol.2020.104362

Zhang, Z. J., Xia, L., Du, J., Li, S. W., and Zhao, F. (2021). Cloning, characterization, and RNAi effect of the chitin synthase B gene in Cnaphalocrocis medinalis. J. Asia-Pac. Entomol. 24(1): 486–492. https://doi.org/10.1016/j.aspen.2021.03.004

Zheng, X., Ren, X., and Su, J. (2011). Insecticide susceptibility of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae) in China. J. Econ. Entomol. 104(2): 653–658. https://doi.org/10.1603/EC10202

Zhou, Y. J., Du, J., Li, S. W., Shakeel, M., Li, J. J., and Meng, X. G. (2021). Cloning, characterization, and RNA interference effect of the UDP-N-acetylglucosamine pyrophosphorylase gene in Cnaphalocrocis medinalis. Genes 12(4): 464. https://doi.org/10.3390/genes12040464

Zhu, P., Zheng, X., Johnson, A. C., Chen, G., Xu, H., Zhang, F., and Gurr, G. M. (2022). Ecological engineering for rice pest suppression in China: A review. Agron. Sustain. Dev. 42(4): 69. https://doi.org/10.1007/s13593-022-00788-5