Abstract
Eggs of the greater wax moth, Galleria mellonella (L.) and the Indian meal moth, Corcyra cephalonica Stalion were evaluated as hosts for mass-rearing Trichogramma pretiosum Riley. The efficacy of the Trichogramma reared on both the hosts were studied on fall armyworm, Spodoptera frugiperda (J.E. Smith). Parasitoids developed from each source were evaluated on the source host and on the target host, S. frugiperda. T. prestiosum reared on the two factitious hosts viz., C. cephalonica and G. mellonella were able to parasitize S. frugiperda eggs under laboratory conditions. Parasitoids reared on G. mellonella (GF) exhibited significantly higher parasitism rate and lower host larval emergence than the parasitoids reared on C. cephalonica (CF) when tested against S. frugiperda. The results demonstrate that the choice of rearing host influences the parasitism efficiency, host suppression, and fitness traits of T. pretiosum. These findings support the development of optimized mass-rearing protocols and the application of this parasitoid in biological control strategies targeting S. frugiperda.
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Introduction
The fall armyworm, Spodoptera frugiperda (J.E. Smith) is a highly destructive and notorious pest which poses a significant threat to agricultural production worldwide. Known for its polyphagous feeding behavior, FAW infests over 350 plant species across 76 plant families, with a preference for maize (Zea mays L.)1 and exhibits a high capacity to establish in a wide range of warm climatic conditions2,3,4. In India, particularly in Tamil Nadu, field studies have shown that FAW infestations can lead to maize yield losses ranging from 70 to 80%, posing a serious economic threat to smallholder farmers5.
The pest is native to tropical and sub-tropical America6. The first emergence of this invasive pest in India was reported in maize fields in Shivamogga district of Karnataka in 2018 and within a span of 1–3 years the pest spread throughout India7. In northeast India, outbreak of FAW was first reported on 2019 from Lunglei district of Mizoram. Its invasion has changed the status of pests in Maize and other important crops8.
Annual losses from FAW on maize ranged from 8.3 to 20.6 million tons9. Although several control measures have been adopted including chemical control, biological control, physical methods to reduce the crop yield loss10; however chemical control is opted mostly considering the faster spread of damage of the pest. Its invasion has substantially increased production costs, primarily due to greater reliance on chemical pesticide applications, placing additional financial strain on farmers9. Although chemical insecticides remain the predominant method of control, their intensive use has raised growing concerns about environmental contamination, development of resistance, pest resurgence, and adverse effects on non-target organisms.
This pest can continue their generations throughout the year due to its strong flying and dispersing capacity aided by high fecundity and polyphagous nature of the pest. Therefore, a sustainable management approach alternative is deployment of egg parasitoids such as T. pretiosum which will control the pest effectively. These parasitoids are effective biocontrol agents, capable of suppressing pest populations at the egg stage, thereby reducing larval emergence and subsequent crop damage11,12,13.
Members of the genus Trichogramma are known to parasitise and suppress populations of at least 28 Lepidopteran pest species affecting agricultural crops, forest ecosystems, and fruit trees14. For effective implementation in biological control programmes, it is essential to screen and mass-release appropriate Trichogramma species, as their performance can vary depending on the host species and environmental conditions11,15,16. The large-scale application of egg parasitoids hinges on the continuous availability of parasitoids, which in turn depends on reliable access to suitable factitious host eggs for mass rearing. Utilizing multiple host species simultaneously in rearing protocols may enhance the stability of parasitoid production. This approach allows for the flexibility to switch parasitism to an alternative host in case of quality deterioration or reduced availability of the primary host species. Maintaining alternative host options in mass-rearing systems can mitigate production disruptions and ensure consistent supply17.
An accurate estimation of host suitability is the key to achieving successful biological control programmes, particularly in optimizing both mass-rearing efficiency and field efficacy. The rice moth, C. cephalonica, is widely used as a factitious host for rearing Trichogramma spp. in several Asian countries, including India, due to its local availability and ease of culture18,19,20. However, parasitoids reared on a single host species are often released to target multiple pest species, which can introduce performance inconsistencies, as host-derived traits may not translate across different targets. An alternative host, the greater wax moth (Galleria mellonella), presents several advantages. Its eggs are readily accepted by a variety of Trichogramma species17,21 and it can be efficiently reared on artificial diets under laboratory conditions. Additionally, female G. mellonella exhibit high fecundity, typically laying between 500 and 1800 eggs per individual, with an average daily output of approximately 60 eggs22,23,24. Haque et al.17. further demonstrated that G. mellonella outperformed Plodia interpunctella (Hübner) as a host in cold-storage-based rearing systems, largely due to its eggs maintaining higher quality and suitability for parasitism following low-temperature exposure, an essential trait for long-term egg storage. Moreover, Stein25 reported that T. minutum reared from G. mellonella eggs under variable temperature and light conditions parasitised more eggs of the codling moth (Laspeyresia pomonella L.) than those reared from Sitotroga cerealella (Olivier) under constant environmental conditions, highlighting the influence of host origin and rearing environment on parasitoid performance.
Given the critical importance of consistent host egg availability for the large-scale production of egg parasitoids, this study aimed to assess the suitability of G. mellonella eggs for mass rearing of T. pretiosum in comparison to the traditionally used host, C. cephalonica, under controlled laboratory conditions. The primary objective was to evaluate and compare the parasitism efficiency and developmental performance of T. pretiosum reared on these two factitious hosts when subsequently deployed against the target pest, S. frugiperda. If both host species support effective parasitism of S. frugiperda eggs, then determining the superior host for mass rearing would depend on key performance indicators. In this context, adult emergence rate, parasitism rate and proportion of female progeny were considered as primary parameters for assessing host suitability. These were further evaluated alongside secondary biological traits, including adult longevity, and total developmental duration, to provide a comprehensive understanding of host influence on parasitoid quality and production potential.
Materials and methods
Insect cultures
The initial cultures of C. cephalonica and T. pretiosum were obtained from the State Biological Control Laboratory, Upper Shillong, Meghalaya, India; and the initial cultures of G. mellonella and S. frugiperda were obtained from the National Bureau of Agricultural Insect Resources (NBAIR), Bengaluru, India.
Establishment and maintenance of insect populations
The insect populations were cultured and maintained separately in the School of Crop Protection Laboratory, CPGSAS, CAU, Umiam, Meghalaya under the controlled conditions at 25 ± 1 °C with 60–70% R.H. and 14:10 h (L: D) throughout the rearing period.
C. cephalonica was reared in Corcyra rearing boxes of size (43 cm×23 cm×12 cm, L×W×H) on an artificial diet consisting of crushed rice grains (2.5 kg), crushed raw groundnut (400 g), yeast powder (10 g), wettable Sulphur 80% (5 g), Streptomycin sulphate (0.5 g) following protocol suggested by Lalitha and Ballal26.
G. mellonella was reared using a slightly modified diet that was developed by Jones et al.27. by using corn meal (400 g), wheat flour (200 g), wheat bran (200 g), milk powder (200 g), yeast (100 g), honey (400 ml) and glycerine (300 ml).
S. frugiperda were maintained on a semi-synthetic diet (kidney beans and wheat-germ based diet) modified from Pinto et al.28.
The adults of the laboratory cultures were then sub-cultured and their population was maintained for the study. Less than 24 h old eggs of all the three insects were used for study in the experiment.
T. pretiosum culture were maintained for 5–6 generations on the two factitious hosts viz., C. cephalonica and G. mellonella prior to the tests. The parasitoids were provided with freshly laid host eggs regularly. Cotton swab with 20% honey solution + vitamin E was provided as a source of food for the adult parasitoids.
Experimental procedure
Four treatments were established to evaluate the parasitoid performance: (1) parasitoids reared on C. cephalonica and tested on C. cephalonica (CC), (2) reared on G. mellonella and tested on G. mellonella (GG), (3) reared on C. cephalonica and tested on S. frugiperda (CF) and (4) reared on G. mellonella and tested on S. frugiperda (GF). Each treatment was replicated five times.
Approximately 30 numbers of eggs (< 24 h old) of C. cephalonica, G. mellonella and S. frugiperda were used separately to prepare egg cards (1 × 2 cm size) separately. Excess eggs and scales were carefully removed using a fine brush under a binocular microscope. Each egg card was placed in glass vials (5 ml) and were exposed to a pre-mated adult female of T. pretiosum for all the treatments. 20% honey solution + vitamin E was smeared on the walls of the vials as food for Trichogramma and the vials were covered to prevent the possible escape of adult parasitoids. The female parasitoids were allowed to oviposit for 24 h and removed thereafter. Exposed eggs were kept separately and allowed to develop under the same conditions used for parasitoid colony maintenance.
Parasitoid efficiency measurements
Four days after the removal of female parasitoids, the number of parasitised (blackened) eggs was recorded using a stereoscopic microscope. Observations continued daily until no further parasitoid emergence was noted, confirming the completion of adult emergence from host eggs. The emergence rate of T. pretiosum was assessed by counting the number of blackened eggs with visible emergence (exit) holes. Adult longevity was determined by isolating newly emerged individuals in glass vials without access to food or water, and monitoring them until death. The proportion of female progeny was calculated by examining fully developed, dead adults under a microscope. The percentage of host larval emergence was recorded by counting the number of larvae hatched from eggs exposed to female parasitoids. Additionally, the percentage of damaged or unhatched eggs was determined based on the total number of eggs presented for parasitization in each treatment.
The parameters were calculated using the equations below:
Statistical analysis
All data were subject to normality test via Shapiro-Wilk test, wherein low p-value of less than 0.05 was observed indicating non-suitability of performing ANOVA. Therefore, the response of the parasitoid to the treatments were analysed by performing paired t-test and comparing the means of treatments at p ≤ 0.05 for all the trials. All the statistical analysis was carried out with the help of R statistics.
Results
Effects of factitious hosts on the parasitism of parasitoid, larval emergence of host and adult emergence of parasitoid
There were significant differences in the per cent parasitization of T. pretiosum with respect to the CF and GF treatments (t = −4.99, p < 0.05) whereas no significant differences were observed in the CC and GG treatments (t = 0.125, p = 0.906) (Table 1). The parasitisation was maximum in GF treatment (60.67%) and minimum in CF treatment (40.00%) (Table 1). Significant differences were observed in the per cent larval emergence of host in the CC and GG treatments (t = −4.42, p < 0.05) whereas no significant differences were observed in the CF and GF treatments (t = 2.622, p = 0.058) (Table 1). Per cent larval emergence of host was lowest on GF treatment which recorded 22.00% (Table 1). Per cent adult emergence of parasitoid were higher than 55% in all the treatments and maximum was observed in GF treatment (Table 1); and were significantly different in CC and GG treatments (t = 3.1, p < 0.05). There were no significant differences in the per cent adult emergence of T. pretiosum with respect to the CF and GF treatments (t = −1.68, p = 0.1673).
Effects of factitious hosts on the female progeny and damaged or unhatched host eggs
There were no significant differences observed in the per cent female progeny of T. pretiosum in CC and GG treatments (t = −1.15, p = 0.31) whereas significant differences were seen with respect to the CF and GF treatments (t = −2.85, p < 0.05) (Table 2). Moreover, the percentage of female progeny was over 55% in all treatments (Table 2). Significant differences were observed in per cent damaged or unhatched host eggs in CF and GF treatments (t = 3.49, p < 0.05) while no significant differences were observed in CC and GG treatments (t = 0.438, p = 0.683) (Table 2). Highest damaged or unhatched host egg was seen on CF treatment (36.00%) (Table 2).
Effects of factitious hosts on total developmental period and adult longevity of parasitoid
There were significant differences in the total developmental period of parasitoid in CC and GG treatments (t = −3.810, p < 0.05); whereas no significant differences were observed in CF and GF treatments (t = 1.633, p = 0.177) (Table 3). The shortest development time was observed in the CC treatment (Table 3). Adult longevity of the parasitoid did not differ significantly across the treatments. However, higher adult longevity of the parasitoid was recorded in CF and GF treatments (Table 3).
Discussions
Our study evaluated the suitability of two factitious hosts viz., C. cephalonica and G. mellonella for mass production of the egg parasitoid, T. pretiosum and recorded the following parameters i.e., parasitization, larval emergence of host, adult emergence of parasitoid, female progeny, damaged or unhatched host eggs, total developmental period and adult longevity of the parasitoid. T. pretiosum females accepted both the tested host eggs for parasitism. Notably, parasitoids reared on G. mellonella (GF) exhibited significantly higher parasitism rate and lower host larval emergence than the parasitoids reared on C. cephalonica (CF) when tested against S. frugiperda. However, significantly higher female progeny was recorded for parasitoids reared on C. cephalonica (CF treatment) than the parasitoids reared on G. mellonella (GF) when tested against S. frugiperda. There were no significant differences in the adult emergence of T. pretiosum with respect to the CF and GF treatments.
In this study, parasitoids reared on both factitious hosts were evaluated against their respective rearing host and a target host, S. frugiperda, to determine their efficacy and adaptability. For these, we evaluated various parameters among which parasitism rate, adult emergence rate and proportion of female progeny were considered as primary parameters for determining host suitability. However, we consider parasitism rate of T. pretiosum as the most important parameter for adoption in biological control as these parasitoids are considered as biotic insecticide i.e., control will be achieved by the inundative release of parasitoid population itself and not by their progeny.
The results indicate that factitious hosts used for mass rearing influence the parasitism and performance of T. pretiosum. Maximum parasitisation was observed in GF treatment (60.67%) and minimum in CF treatment (40.00%). Earlier findings of differences in parasitism efficiency were reported in T. minutum29; T. evanescens30; and T. pretiosum31 when reared on various host species. Significant differences in the per cent parasitization of T. pretiosum with respect to the CF and GF treatments were observed where the parasitisation was more in GF compared to CF treatment. Haque et al.17. also reported higher parasitism rates of T. evanescens in G. mellonella eggs compared to Plodia interpunctella. Montoya32 recorded 45–55% egg parasitism of Trichogramma spp. in the maize fields in Mexico. While, Peralta et al.33. reported average parasitism of less than 15% by Trichogramma sp. on FAW eggs in Tamaulipas, Mexico.
The sex ratio across all the treatments was female-biased, with a relatively high proportion of female progeny observed. Significantly higher percentage of female emergence (68.52%) was recorded in CF than GF treatment. These findings are in line with those of El-Wakeil30who reported a female-biased sex ratio in T. evanescens, ranging from 63.6 to 74.4% across various host treatments. Similarly, Rathi and Ram34 found higher female emergence in T. chilonis reared on Achaea moorei (74.90%), Earias vittella (73.30%), H. armigera (71.50%), and C. cephalonica (68.70%). Similarly, Dileep35 recorded 60% female population of T. chilonis from the eggs of C. cephalonica and T. pretiosum yielded a higher proportion of females from S. frugiperda eggs (sex ratio of 1:2.1) compared to those reared on C. cephalonica (1:1.53). These variations suggest that host species and associated factors—such as egg quality, nutritional content, and maternal effects- can influence sex ratio outcomes. Thus, selection of an appropriate factitious host for mass rearing must consider not only parasitism efficiency and emergence but also potential impacts on progeny sex ratio.
Adult longevity of T. pretiosum is known to be influenced by the host species, with previous studies reporting that females typically survive for only 2–3 days in the absence of food30. In the present study, however, no significant differences in adult longevity were observed across the different host treatments. This lack of variation may be attributed to the controlled and consistent laboratory conditions, which could have minimized potential effects of host species on longevity. Butler and Lopez36 similarly reported slightly greater longevity of T. evanescens reared on Helicoverpa eggs compared to those reared on S. cerealella, attributing such differences to the nutritional composition of the host eggs. These findings suggest that although host quality can influence parasitoid lifespan, its effects may be masked under standardized environmental conditions, emphasizing the need to evaluate such traits under more variable or field-relevant settings.
Parasitism efficiency and parasitoid fitness are commonly assessed through parameters such as percent parasitism, adult emergence rate, sex ratio, and developmental duration37,38,39,40,41,42,43. In case of our findings where GF treatment resulted in better performance of parasitism rate over CF treatment; whereas CF treatment resulted higher female progeny than GF treatment. Moreover, considering the parasitoid’s adult emergence, female progeny, longevity, damaged or unhatched host egg it can be inferred that G. mellonella used as factitious host for mass rearing of T. pretiosum will result in better control of S. frugiperda. Having considered all these parameters, it can be suggested that G. mellonella will serve as better potential hosts for mass multiplication of T. pretiosum for managing S. frugiperda in biocontrol.
Abdel-Galil et al.44. validated whether the artificial or natural diets are the best for G. mellonella and P. interpunctella hosts for rearing Trichogramma turkestanica Meyer and reported G. mellonella reared on artificial diets to be the best host with successful parasitized eggs, emerged adult parasitoids from parasitized eggs, and emerged female parasitoids as compared to P. interpunctella. In a prior evaluation of C. cephalonica and G. mellonella as hosts for the larval ectoparasitoid Habrobracon hebetor (Say), Haque et al.17. identified G. mellonella as the superior host, largely due to its larger larval size and extended developmental period.
Host quality and developmental rate can significantly affect parasitoid biology, influencing key traits such as adult size, fecundity, and longevity45. Furthermore, morphological and structural differences in host eggs may also affect their overall suitability for parasitism, contributing to variation in developmental success.
While this study demonstrates the potential of T. pretiosum for parasitizing S. frugiperda using factitious hosts under controlled laboratory conditions, it is important to recognize that these results reflect a primary screening under artificial settings. The true effectiveness and ecological adaptability of the parasitoids must be validated under natural field conditions. In such environments, various biotic and abiotic factors—including crop architecture, plant morphology, prey distribution, and climatic variability—can significantly influence the foraging behavior, dispersal, and overall efficiency of natural enemies46,47,48,49,50,51,52. Specifically for Trichogramma spp., parasitism rates are known to vary across different habitats, plant species, and plant structures. Factors such as plant spacing, volatile compounds, and chemical profiles may all play roles in modulating the parasitoid’s host-seeking behavior and parasitism efficiency53,54. Therefore, further field-based investigations are essential to confirm the applicability of these findings and to optimize the use of T. pretiosum in integrated pest management programs.
Other important factors which need to be addressed for suitable host for mass multiplication are production cost and field efficacy. 70% of space and cost is related to the factitious host production for mass production of Trichogramma parasitoid55. Rearing techniques also influence parasitoid quality (parasitism, fecundity, searching capacity etc.) impacting effectiveness of parasitoid in field release while field efficacy is affected by appropriate time of parasitoid release and sufficient number of releases etc.56. Continuous availability of host for cost effective mass production of T. pretiosum is an important factor for successful biological control, therefore, further investigations on other criterias like rearing techniques for mass multiplication, production costs, field efficacy, is necessary.
Conclusions
Assessing fitness parameters in egg parasitoids is essential in both behavioral ecology and biological control research. In this study, T. pretiosum reared on two factitious hosts, C. cephalonica and G. mellonella, successfully parasitised eggs of S. frugiperda. While both hosts supported parasitoid development, G. mellonella demonstrated several advantageous traits that may favor its use in mass rearing. Moreover, host availability and rearing feasibility further support its potential as a preferred host species. These findings suggest that G. mellonella can be considered a viable alternative to C. cephalonica for large-scale production of T. pretiosum. However, additional field evaluations are necessary to validate its effectiveness under natural conditions and to optimize its integration into biological control programs targeting S. frugiperda.
Data availability
All the supporting data of this study are available with the manuscript. Supplementary information and additional data will be made available from the corresponding author upon reasonable request.
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Acknowledgements
The authors are thankful to the College of Post Graduate Studies in Agricultural Sciences, Central Agricultural University (Imphal); State Biological Control Laboratory, Upper Shillong and National Bureau of Agricultural Insect Resources, Bengaluru for providing necessary facilities, constant support, encouragement and cultures of insects for conducting the research.
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K.N. planned the experiment, methodology, conceptualization, supervision, visualization and contributed in writing the manuscript. Z.M. conducted the experiments, collected data and preparation of manuscript. M.P. contributed to analysis and arrangement of the manuscript. J.G. contributed to the statistical analysis of data and writing of manuscript. M.S. contributed to the statistical analysis of data, editing and correction of the manuscript.
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Ningthoujam, K., Moirangthem, Z., Pathak, M. et al. Host suitability and parasitism of Trichogramma pretiosum on Galleria melonella and Corcyra cephalonica for the management of fall armyworm. Sci Rep 15, 34154 (2025). https://doi.org/10.1038/s41598-025-14656-y
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DOI: https://doi.org/10.1038/s41598-025-14656-y
