Oviposition behavior and larval attraction of the fall armyworm Spodoptera frugiperda to different maize plant varieties

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Akinbuluma, Olubisi O. Bamifewe, Olajumoke Y. Alabi, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4601270/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Phytophagous insects likely select suitable host plants for oviposition based on olfactory and tactile cues. However, details of how insects differentiate among different plant varieties are often unclear. The fall armyworm ( Spodoptera frugiperda J. E. Smith) is a highly destructive pest on maize, but little is known about the attraction and oviposition preference of S. frugiperda to different maize varieties, particularly in the context of sub-Saharan Africa, where the insect is a major threat to maize production. We determined the oviposition preference of S. frugiperda females on six different maize plant varieties three of which were hybrid varieties and three were open pollinated varieties, in multiple-choice and no-choice assays. We also evaluated the attraction preference of S. frugiperda larvae on these maize varieties, using an olfactometer bioassay. We found that S. frugiperda females oviposited significantly less egg masses on the hybrid varieties DEKAIB and 30Y87 than on the other varieties tested, and that females oviposited less on the hybrid maize varieties compared to the open pollinated maize varieties overall. Additionally, we found that S. frugiperda larvae were more attracted to the open pollinated variety LMFP than to clean air, which was not the case for any of the other maize varieties tested. Taken together, our results show that S. frugiperda responds differentially to the different maize varieties and that hybrid maize varieties seem less attractive. Further investigating the chemistry of hybrid maize varieties like DEKAIB might yield clues on how to breed maize varieties with increased resistance against S. frugiperda infestation. Fall armyworm maize plant volatiles multiple choice experiment olfactometer bioassay resistant varieties Figures Figure 1 Figure 2 Figure 3 Introduction The use of improved crop varieties is one of the key practices towards an efficient integrated pest management programme against herbivorous pests and pathogens (Acquaah 2004 ; Liebman and Davis 2009 ). One focus of plant breeding programmes is generally to obtain hybrids that are able to withstand extreme conditions of cold or drought or that have resistance to pathogenic organisms or insect pests (Kutka 2011 ; Lee et al. 2015 ; Kumar et al. 2020 ; Enders and Begcy 2021 ). Maize ( Zea mays L.) is a versatile multi-purpose crop, used as a feed globally and important food crop, especially in sub-Saharan Africa and Latin America (Erenstein et al. 2022). It is also one of the most important cereal crops in Nigeria grown for human consumption, animal feed and several industrial uses (Iken and Amusa 2004 ; Abdulraham and Kolawole 2006 ; Odeyemi et al. 2020 ). Nigeria is the largest producer of maize in Africa with a production of 10.2 million metric tons on 4.8 million hectares of land, representing 0.42% of the total global production (FAO, 2018; Kamara et al. 2020). In maize, open-pollinated (OP) varieties are traditionally used varieties that inter-pollinate freely during seed production, resulting in heterogeneous varieties. The OP varieties have broad genetic bases selected by the environment and farmers over many generations, which help to maintain moderate stress resistance and yield characteristics (Dávila-Flores et al. 2013; Lima et al. 2022). Thus, OP varieties are valuable genetic resources of breeding programmes (Costa et al. 2018 ; Costa et al. 2020 ; Dos Santos et al. 2020 ; Sing et al. 2021; Lima et al. 2022). In contrast to OP varieties, hybrid maize varieties are cultivated varieties that result from the fertilization of one maize plant by another genetically unrelated plant through controlled cross-pollination (FAO 2016 ). The quality of hybrid maize depends greatly on methods of field production, both in adherence to quality assurance standards and implementation of appropriate agronomic management (Bedő and Barnabás 2013 ; MacRobert et al. 2014 ; Karim et al. 2018 ). Maize hybrid seeds provide farmers with varieties that have improved specific traits, such as high yield potential and unique trait combinations to counter diseases and adverse growing conditions (MacRobert et al. 2014 ), for example increased resistance to insect attack (Barry et al. 1992 ; de Lange et al. 2014 ) and pathogens (Wang et al. 2008 ). The fall armyworm, Spodoptera frugiperda (J.E. Smith) is a major phytophagous pest of agricultural crops endemic to tropical and subtropical regions of the Americas (Sparks, 1979 ; Johnson et al. 1987; Nagoshi et al. 2012 ). It is a migrant pest with a wide host range causing great economic loss whenever present. The pest can feed on over 350 host plants belonging to 76 families, with the Poaceae being the most preferred host (Montezano et al., 2018 ). Since S. frugiperda invaded Africa (Georgen et al. 2016), it was reported in more than 45 African countries, affecting maize production (Kasoma et al. 2021 ; Edosa et al. 2021) with over 50% of maize field losses (De Groote et al. 2020 ; Abro et al. 2021 ). In Nigeria, S. frugiperda has established itself in three agroecological zones, causing extensive damage to maize farms, particularly in the humid forest agroecological zone in southwestern Nigeria (Ojumoola and Omoloye 2023 ), causing between 50–80% damage in many maize fields (Odeyemi et al. 2020 ). One potential approach to S. frugiperda management is to explore the role of maize varieties in influencing the attraction and acceptance behaviour of S. frugiperda . Host plants play an important role in the chemical ecology and behaviour of S. frugiperda , because the insects respond to chemical cues for oviposition and larval feeding (Guo et al. 2021 ; Sotelo-Cardona et al. 2021; Zhang et al. 2023 ). In general, different maize varieties, whether hybrid or OP, vary in their biochemical composition, which make them more or less suitable to S. frugiperda (Yang et al. 2023 ). For example, some maize varieties produce higher levels of secondary metabolites such as terpenoids, which have been shown to have insecticidal properties, while other varieties may produce compounds that are attractive to S. frugiperda (Firake et al. 2020). In addition, physical characteristics of host plants, such as trichomes, wax amount, thickness and toughness of leaves and secondary toxic metabolites influence host-plant selection behavior (Gatehouse 2002 ). Differential preference of S. frugiperda was reported in maize cultivars due to difference in cuticular lipids (Yang 1993a), presence of wax materials on the leaf surface (Yang 1993b) and anti-feedant and anti-repellent properties (Tiwari 2022 ). Several studies have investigated the oviposition preference of S. frugiperda to identify which host plant species are either resistant or susceptible to the insect pest (Ba et al. 2020 ; Guo et al. 2021 ; Tiwari 2022 ; Sisay et al. 2023 ). Only few studies evaluated oviposition preference of S. frugiperda on different maize varieties (He et al. 2021 ; Zhang et al. 2023 ). The use of maize varieties that are less preferred by S. frugiperda could reduce the level of infestation in the field. Whether Nigerian S. frugiperda females show any oviposition preference towards specific hybrid or OP maize varieties is largely unknown. In addition, little is known about the larval attraction of S. frugiperda to different maize plant varieties, particularly in Nigeria (Odeyemi et al. 2020 ). To evaluate whether S. frugiperda show differential attraction and oviposition behaviours towards different maize varieties, we investigated the oviposition preference of the Nigerian S. frugiperda population on six maize varieties, three of which were hybrid and three were OP varieties. To determine whether the number of egg masses laid by S. frugiperda female increased or decreased over the oviposition period, we checked for variability in the timing of egg-laying on the maize varieties, by conducting oviposition assay over three days in both multiple choice and no-choice experiments. Finally, we determined whether the S. frugiperda larvae were attracted differentially towards the odors from the different maize plant varieties. Materials and Methods Study location The experiment was conducted in the Entomology Research Laboratory and Insect Chemical Ecology Laboratory, Department of Crop Protection and Environmental Biology, University of Ibadan, Nigeria, under ambient conditions of 27 ± 1°C and 65 ± 5% RH. Source of maize seeds and planting Three hybrid maize varieties (referred to in italics in the remainder of this manuscript), DEKAIB, 30Y87 and P3966W , and three OP varieties (referred to in bold), LMFP, SWAN1 and V9928 , were used for experiments. DEKALB was obtained from International Institute of Tropical Agriculture; V9928, LMFP and SWAN1 were obtained from Institute of Agriculture Research and Training, Moor Plantation, Apata Ibadan, while 30Y87 and P3966W were obtained from Corteva Agriscience. These varieties were selected because they are cultivated by Nigerian farmers, but so far were never tested for attraction and oviposition by S. frugiperda . Two seeds from each variety were sown in a 10 kg pot filled with heat-sterilized loamy soil. The potted plants were grown and maintained in a green house at 26 ± 2°C, 70 ± 5% RH and a photoperiod of 14:10 (L:D) and each variety was planted in six pots. To have a sufficient number of plants for the experiments, the plants were sown every week. Seven days after each sowing, maize plants in each pot were thinned to one. For the experiments, 14-day old potted maize plants were used. Insect collection and rearing Spodoptera frugiperda larvae were collected from a naturally infested maize farms at Sasha-Ajibode (latitude 7 o 28 1 37.70688 E, 3 0 54 1 N) and Elekuru-Akinyele areas (7 0 36 1 25.54092,3 0 49 1 N) of Ibadan, Southwestern Nigeria and reared at the Entomology Research Laboratory, University of Ibadan, Nigeria. The larvae were separated into a transparent sauce cup (EEZEE, Nigeria; 4 cm ×3 cm; 40 mL) with one larva per cup, which were fed daily with fresh maize leaves of the SWAN 1 variety and covered with lid until pupation. The pupae were placed into clean vials lined with moist tissue paper until adult emergence. The adult insects were kept in the same vials and fed with 10% sugar solution. For mating to occur, 2–3 day old virgin adults were paired in mating cups (AVT Plastics, 500 mL, one pair/cup) with a sauce cup (4 cm ×3 cm; 40 mL) filled with cotton wool soaked in a 10% sugar solution and covered with muslin cloth. Eggs were collected daily and newly emerged larvae were individually placed in cups and fed with fresh maize leaves of the SWAN 1 variety (approx. 1 g/cup) for 7–12 days. For the experiments, 2–4 day-old mated adult females and the 7–12 day old larvae of the new generations were used. Oviposition assays Oviposition preference and performance of S. frugiperda females was determined in multiple-choice and no-choice experiments inside cages enclosed in the same green house as described above, using intact plants from the six maize varieties. In the multiple choice test, six pots were properly labeled and placed in a mesh cage (40 × 35 × 50 cm), whereby each pot contained one 14 day-old plant of one maize variety. The potted plants were arranged inside the cage in a completely randomized design (CRD). One 2–4 day-old mated female was released into the cage for 72 hours. The multiple choice test was conducted in four replicates. Each replicate contained different batch of the six plants (of the same age) and was set up separately at one week interval. The number of egg masses on the maize plants inside the cage was counted and recorded every 24 h for a period of 72 h. In the no-choice experiment, we placed one pot containing one 14 day-old plant of each variety within similar mesh cages (40 × 35 × 50 cm) together with one 2–4 day-old mated female. The no-choice experiment thus had six treatments with 4 replications for each variety. The number of egg masses on the maize plants was counted and recorded every 24 h for a period of 72 h. Egg masses were carefully counted and recorded from outside the cage such that female insects were not disturbed. All data were collected between 13:00–16:00 daily and the bioassays were conducted between March and June 2022. Responses of Spodoptera frugiperda larvae to intact plants To determine the orientation responses of S. frugiperda larvae to intact maize plants, we chose to test the two most preferred varieties and two least preferred varieties based on the oviposition preference of S. frugiperda females from the experiment above. The experiment was conducted in a Y-tube olfactometer, as described in Akinbuluma and Chinaka ( 2023 ) with some modifications, and summarized here. The experimental arena consisted of a horizontal pyrex glass Y-tube (10 mm i.d; stem 85 mm; arms 75 mm at a 60° angle to the stem). Air from a field pump was passed through activated charcoal and humidified with double distilled water. The airflow was split into two halves. One half was passed through a glass chamber with a tightly sealed lid, enclosing a potted plant (test) and into one arm of the olfactometer, while the other half of the airflow was passed through an empty glass chamber (control) at the same regulated flow rate of 60 mL/min. A vacuum line was connected to a mini pump powered by a rechargeable battery pulled air through the two arms of the Y-tube. Fourth instar S. frugiperda larvae were placed in the bioassay room for at least 12 h before the experiment to acclimatize them to the room conditions. The larvae were also starved for 3 hours before the experiment by removing the larvae from their feed container to a small empty rearing cup (1 larva/cup). Twelve larvae were individually assayed in random order on the plant varieties such that 1 larva was assessed 4 times on the four maize varieties. Larval choice was recorded after spending 3 mins in any of the arms of the Y-tube. Insects that failed to choose an arm within 10 mins were recorded as non-responders and were not included in analysis. After testing on each maize variety, the olfactometer set-up, glass chambers and connections were wiped with 70% ethanol and dried to avoid odor contamination between consecutive bioassays and the odor source positions were also exchanged. After an interval of 15 mins, the next larva was tested. Bioassays were always conducted between 13:00 and 18:00 hours. This study was conducted between May and July 2022. Statistics and Data Analysis The multiple-choice and no-choice oviposition experiments were analyzed separately. The number of egg masses was modeled with generalized linear models (GLM, McCullagh and Nelder, 1989 ) using Poisson regression. Both models contained maize variety and duration of oviposition as well as their interaction as explanatory variables. In addition, the position of the egg masses in the multiple choice experiment was determined using a separate model, containing position, hours and their interaction. Finally we investigated if there were differences between the Hybrid varieties ( DEKAIB, 30Y87 and P3966W ) and OP varieties ( LMFP, SWAN1 and V9928 ) with a model that contained the varietal type and hour, as well as their interaction as explanatory variables (Table 1 ). Posthoc comparisons and testing the trends of egg laying over time were done on marginal means (Lenth 2023 ) and the result displayed with the compact letter display (Piepho, 2004 ), with p = 0.05 as significance level and Tukey adjustment for multiple testing. The olfactometer experiment data were analyzed using chi-square tests. All statistical analyses were performed in R (R Core Team 2022 ). Table 1 Overview of the explanatory factors in models used. In all models, the response was the number of egg masses laid by female Spodoptera frugiperda Model Model (Explanatory variables) N (cases) Df Choice test Variety * hour 72 64 Position of egg masses Position * hour 84 78 No-choice test Variety * hour 72 64 Maize-type test Type * hour 72 54 Results Oviposition preference of adult S. frugiperda female When we checked the number of egg masses laid by S. frugiperda female on the different maize varieties in the multiple choice experiment, we found significant difference between the maize varieties (p < 0.05, n = 72, df = 64), but no significant interaction between hours of oviposition and maize variety. Significantly fewer egg masses were laid over the 72-hour period on P3966W and LMFP but not on DEKAIB, 30Y87 , SWAN 1 and V9928 (Fig. 1 A). However, after 24 h of oviposition, the number of egg masses found on DEKAIB was significantly lower than those of P3966W , LMFP and V9928 varieties (Fig. 1 A). Similarly, in the no-choice experiment, we found significant difference between the maize varieties but no interaction with hours of oviposition (p < 0.05, n = 72, df = 64). Across 72 h oviposition period, we observed significantly fewer egg masses (p < 0.05) of S. frugiperda on maize varieties, LMFP and SWAN 1 , but not on DEKAIB, 30Y87 and P3966W (Fig. 1 B). However, there was no significant difference in the number of egg masses laid across all maize varieties after 24 h of oviposition (Fig. 1 B). When we combined the egg masses over time for each maize variety in the multiple choice experiment (Fig. 1 B), we found that DEKAIB and 30Y87 had significantly lower egg mass numbers than varieties, LMFP, SWAN 1 and V9928 (p 0.05). Similarly, in the no-choice experiment, we found significantly lower egg mass number on DEKAIB and 30Y87 than on LMFP maize varieties (p < 0.05) (Fig. 1 D). When combining all three hybrid varieties and the OP varieties, egg mass counts only significantly differed between 24 h and 72 h in the multiple choice (Fig. 2 A), with no significant difference in the number of eggs laid over 72 h in the no-choice experiment (Fig. 2 A and B). In the OP varieties, we found significantly lower egg masses on the hybrid varieties after 72 h of oviposition than after 24 h both in the multiple choice and the no-choice test (p < 0.05). After 24 h of oviposition, significantly fewer egg masses were laid by S. frugiperda female on hybrid maize than on OP varieties in both experiments (p < 0.05) (Fig. 2 A and B). When the cumulative egg mass counts were compared, egg masses on the hybrid varieties were significantly lower than those on the OP varieties in both multiple choice and no-choice experiments (p < 0.05) (Fig. 2 C, D). Response of Spodoptera frugiperda larvae to intact maize varieties Olfactometer experiment with intact plants of the four maize varieties showed that significantly more S. frugiperda larvae (χ 2 = 8.33; p 0.05), 30Y87 (χ 2 = 0.33; p > 0.05) and DEKAIB (χ 2 = 0.33; p > 0.05). Figure 1 Mean number of egg masses (± SEM, n = 4) of Spodoptera frugiperda female. Line plots show number of egg masses on DEKAIB, 30Y87, P3966W (hybrid maize varieties), LMFP, SWAN 1 and V9928 (open pollinated varieties) over 72 h period on maize varieties in (A) multiple choice experiment and (B) no-choice experiments. * = significant difference between mean egg masses over 72 h of oviposition; ns = no significant differences between egg masses between the three time points (24, 48 and 72 h) of oviposition. Different letters under the error bars in a and b show that mean egg numbers between varieties are significantly different after 24h in multiple choice and no-choice experiments, respectively. C) Cumulative number of egg masses after 72 h of oviposition in multiple choice experiment and (D) cumulative number of egg masses in no-choice experiments. Boxes represent the lower and upper quartiles of egg masses on each maize, whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in c and d indicate significant differences among maize varieties (p < 0.05) Figure 2 Mean number of egg masses (± SEM, n = 4 ) of Spodoptera frugiperda female laid on two types of maize varieties. Line plots show the number of egg masses on hybrid and OP maize varieties over a 72 h period in (A) multiple choice experiment and (B) no-choice experiment. Different letters on line show significant difference between mean egg masses laid over time. * in between the error bars in A and B show that mean egg numbers are significantly different after 24 h between hybrid and OP maize varieties in multiple choice (A) and no-choice experiment (B), respectively. (C) Mean cumulative number of egg masses over 72 h oviposition period in the multiple choice experiment and (D) in the no-choice experiment. Boxes represent the lower and upper quartiles of egg masses on each maize type; whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in C and D indicate significant differences between hybrid and OP varieties (p < 0.05) Figure 3 Orientation responses of Spodoptera frugiperda larvae to odors from intact plants and clean air. Bar charts show the preferences of S. frugiperda larvae to odors from intact four maize varieties and clean air presented in a Y-tube olfactometer. Hybrid maize varieties ( DEKAIB, 30Y87 ) are in italics, Open pollinated maize varieties ( V9928, LMFP ) are in bold. Significant means were separated using Chi-square (χ2) test (p < 0.05, n = 12), ns = not significant. Discussion In this study, we investigated the oviposition and attraction of S. frugiperda to different maize varieties, belonging to hybrid and OP varieties, and found i) variation in the number of egg masses laid on maize varieties at different time periods, ii) ovipositional preferences of S. frugiperda females for SWAN 1, V9928 and LMFP compared to DEKAIB and 30Y87 in both multiple-choice and no-choice experiments, iii) significant attraction responses of S. frugiperda larvae towards LMFP maize variety compared to clean air, while this was not the case for V9928 , DEKAIB and 30Y87 . Overall, our results show that DEKAIB and 30Y87 were less accepted and less attractive to S. frugiperda than the other maize varieties. Our results showed significant variation in the number of egg masses laid by S. frugiperda females on the different maize varieties, both in the multiple choice and no-choice tests. The consistently significant lower number of egg masses on DEKAIB suggests that the variety was the least acceptable for oviposition compared to all other tested varieties. Differential oviposition preference of S. frugiperda female for some maize varieties has been shown by others as well (He et al. 2021 ; Zhang et al. 2023 ) and could be due to differences in plant nutrients in maize host plants, which is important for development of offspring (Tiwari 2022 ; Yang et al. 2023 ). For a proper timing of management of S. frugiperda eggs on the maize host plants, it is important to know when egg masses are laid and how this changes over time. Spodoptera frugiperda females have been found to lay reduced numbers of egg batches over time (Russianzi et al. 2021 ). Therefore, we counted the number of egg masses over a three-day period. Among the varieties tested, we found significant reductions in the number of egg masses laid over time. For example, in the no-choice test, significantly more egg batches were laid on the LFMP and SWAN1 maize varieties after 24 h than after 72 h. Also, in the first 24 h, significantly less egg batches were laid on the hybrid maize varieties (Fig. 2 ). We also showed that significantly less egg masses were found on the combined hybrid maize varieties (DEKAIB, 30Y87 and P3966W ) than on the OP varieties ( LMFP, V9928 and SWAN1 ). Even when considered separately, the least number of egg masses were laid on the hybrid varieties compared to the OP varieties (Fig. 1 ). Our results thus indicate that hybrid maize varieties are less favourable to S. frugiperda for oviposition than OP varieties. An earlier report also showed that S. frugiperda did not infest hybrid maize varieties as much as OP varieties in both the lowland and high-altitude land in Kenya (Mutyambai et al. 2022 ). However, Zhang et al. ( 2023 ) found significant oviposition preference of S. frugiperda females for some hybrid maize varieties compared to OP varieties. Thus, differential oviposition cannot always be explained by maize pollination type. Nevertheless, to reduce infestation of maize by S. frugiperda in Nigeria, our results suggest that farmers should use hybrid maize varieties instead of OP varieties for establishing maize fields. We would like to point out that in all our oviposition experiments, we found more S. frugiperda egg masses on the cage surfaces than the maize varieties (Supplementary Fig. SI). This is a common phenomenon that other studies also found (Rojas et al. 2003 ; Barros et al. 2010 ; Meagher et al. 2011 ; Guo et al. 2021 ; Sotelo-Cardona et al. 2021; Volp et al. 2022 ). This may suggest that some maize plants volatiles have repellent effects for female insects, preventing them from ovipositing on the plants. Rojas et al. ( 2003 ) and Barros et al. ( 2010 ) reported that S. frugiperda female oviposited more on corrugated surfaces rather than surfaces treated with some host plant extracts. Thus, our study corroborates the finding that S. frugiperda females lay more eggs on oviposition cage walls away from nearby plants than on plants, which could indicate repellent effects of some plant-derived volatiles (Sisay et al. 2023 ). The olfactometer bioassay with S. frugiperda larvae showed only a larval preference towards LMFP maize variety relative to clean air, and not to the other varieties. Possibly, this variety releases volatiles that are attractive for larvae. Although lepidopteran larvae possess limited mobility, they can still move between plants by way of so-called ‘ballooning’ (e.g. Sokame et al., 2020 ), and also orient towards or away from host plants (Liu et al. 2022). Background volatile compounds have been found to modify larval behavior in S. frugiperda on some maize varieties (Yactayo-Chang et al. 2021 ). Interestingly, S. frugiperda larvae were not significantly attracted to the hybrid DEKAIB and 30Y87 , on which S. frugiperda females oviposited least. In conclusion, our study showed that different maize varieties elicit differential oviposition and larval responses in S. frugiperda. Possibly, variation in maize host plant volatiles may partly account for these different responses. Identification of plant volatiles that attract S. frugiperda females could help to reduce the population. Since the use of plant varieties can affect the extent of infestation and damage by insect pest, varieties with less preference for oviposition by the insect will suffer less infestation and damage compared to those with full acceptance for oviposition by the insect. Our study suggests that planting DEKAIB and 30Y87 , which are hybrid maize varieties could reduce the level of Spodoptera frugiperda infestation in Nigerian fields. Exploring these avenues further will help to develop integrated pest management (IPM) strategies of this pest. Declarations Conflicts of interest/Competing interests The authors declare that they have no conflict of interest/competing interests Supplementary Information Supplementary information can be found on the online version of this article. Availability of data and materials The authors confirm that data supporting the findings of this study are available from the corresponding author on request. Funding: This project was part of the funded project by a grant from the International Foundation for Science, co-sponsored by the Organisation for the Prohibition of Chemical Weapons (OPCW). Author Contribution M.D.A planned and conceived the study and acquired funding. MDA, OOB and MIJT conducted the experiments. OYA contributed towards olfactometer setup. MDA and PR performed data analysis. MDA and OOB wrote the draft manuscript. MDA, PR and ATG reviewed the manuscript. All authors read and approved the manuscript. Acknowledgement We are grateful to the International Foundation for Science, co-sponsored by the Organisation for the Prohibition of Chemical Weapons (OPCW), for funding this work. We thank Dr Felicia O. Aigbokhan of the Corteva Agrisciences, for facilitating the provision of some hybrid maize seeds for this research. References Abdulraham AA, Kolawole OM (2006) Traditional preparation and uses of maize in Nigeria. Ethnobotany Leaflets 10: 291–227. Abro Z, Kimathi E, De Groote H, Tefera T, Sevgan S, Niassy S, Kassie M (2021) Socioeconomic and health impacts of fall armyworm in Ethiopia PLoS ONE, 16, e0257736. Acquaah G (2004) Principles of Crop Production: Theory, Techniques, and Technology. Pearson Education, Prentice Hall: Upper Saddle River, NJ, USA. Akinbuluma MD, Chinaka OP (2023) Efficacy of the parasitic wasp, Dinarmus basalis Rondani (Hymenoptera: Pteromalidae), in reducing infestations by the cowpea beetle, Callosobruchus maculatus (L.) (Coleoptera: Chrysomelidae: Bruchinae). Egypt J Biol Pest Control 33: 1–7. https://doi.org/10.1186/s41938-023-00692-1 Ba TX, Zhang YH, Zhang Z, Guan DD, Li CC, Ji ZY, Yin XT, Zhang AH, Tang QB, Liu YH, Li XR, Zhou X (2020) The host preference and population life tables of Spodoptera frugiperda (Lepidoptera: Noctuidae) fed on maize and wheat. Plant Prot 46:17–23. Barros EM, Torres JB, Ruberson JR, Oliveira MD (2010) Development of Spodoptera frugiperda on different hosts and damage to reproductive structures in cotton. Entomol Exp Appl 137: 237–245. Barry D, Widstrom NW, Darrah LL, McMillian WW, Riley TJ, Scott GE, Lillehoj EB (1992) Maize ear damage by insects in relation to genotype and aflatoxin contamination in preharvest maize grain. J Econ Entomol 85: 2492–2495. Bedő Z, Barnabás B (2013) 60 Years of Hungarian Hybrid Maize 1953–2013, Pannonian Plant Biotechnology Association, Martonvásár, Hungary. pp. 135. Carroll MJ, Schmelz EA, Meagher RL, Teal PE (2006) Attraction of Spodoptera frugiperda larvae to volatiles from herbivore-damaged maize seedlings. J Chem Ecol 32: 1911–1924. Chimweta M, Nyakudya IW, Jimu L., Bray Mashingaidze A (2020). Fall armyworm [ Spodoptera frugiperda (J. E Smith)] damage in maize: management options for food-recession cropping smallholder farmers Int J Pest Manag 66:142–154. Costa EN, Fernandes MG, Medeiros PH, Evangelista BMD (2020) Resistance of Maize Landraces from Brazil to Fall Armyworm (Lepidoptera: Noctuidae) in the Winter and Summer Seasons. Bragantia 79: 377–386. Costa EN, Nogueira L, De Souza BHS, Ribeiro ZA, Louvandini H, Zukoff SN, Júnior ALB (2018) Characterization of Antibiosis to Diabrotica speciosa (Coleoptera: Chrysomelidae) in Brazilian Maize Landraces. J Econ Entomol 111: 454–462. Crabb R (1992) The Hybrid Corn-Makers. West Chicago Publishing Co. Chicago, IL, USA. Day R, Abrahams P, Bateman M, Beale T, Clottey V, Cock M, Witt A (2017) Fall armyworm: impacts and implications for Africa. Outlooks on Pest Manage 28: 196–201. De Groote H, Kimenju SC, Munyua B, Palmas S, Kassie M, Bruce A (2020). Spread and impact of fall armyworm ( Spodoptera frugiperda J. E. Smith) in maize production areas of Kenya. Agriculture, Ecosystem and Environment, 292, 106804. de Lange ES, Balmer D, Mauch-Mani B, Turlings TCJ (2014) Insect and pathogen attack and resistance in maize and its wild ancestors, the teosintes. New Phytol 204: 329–341. Despland E (2021) Selection Forces Driving Herding of Herbivorous Insect Larvae. Front Ecol Evol 9:760806. doi: 10.3389/fevo.2021.760806 . Dos Santos LFC, Ruiz-Sánchez E, Andueza-Noh RH, Garruña-Hernández R, Latournerie-Moreno L, Mijangos-Cortés JO (2020) Leaf Damage by Spodoptera frugiperda J. E. Smith (Lepidoptera: Noctuidae) and Its Relation to Leaf Morphological Traits in Maize Landraces and Commercial Cultivars. J Plant Dis Protect 127: 103–109. Duvick DN, Smith JSC, Cooper M. (2004) Changes in performance, parentage, and genetic diversity of successful corn hybrids 1930–2000. In: Smith CW, Betran J, Runge ECA (ed) Corn: Origin, History, Technology and Production, John Wiley and Sons, Hoboken, NJ, USA, pp 65–97. Edosa TT, Dinka TD (2021) Current and future potential distribution, risk and management of Spodoptera frugiperda . J Innov Agric 8: 14–23. Enders L, Begcy K. (2021) Unconventional routes to developing insect-resistant crops. Mol Plant 14: 439–1453. Erenstein O, Jaleta M, Sonder K, Mottaleb K, Prasanna BM. (2002) Global maize production, consumption and trade: trends and R&D implications. Food Secur 14: 1295–1319. FAO (2016) Seed Security Assessment: Handout 0 - Session 3 (Seed systems). Pages 1–3. http://www.fao.org/agriculture/crops/core-themes/theme/seeds-pgr/know_res/en/ Firake DM, Behere GT (2020) Bioecological attributes and physiological indices of invasive fall armyworm, Spodoptera frugiperda (J.E. Smith) infesting ginger ( Zingiber officinale Roscoe) plants in India. Crop Prot 137: 105–233. Gatehouse JA (2002) Plant resistance towards insect herbivores: A dynamic interaction. New Phytol 156:145–169. Goergen G, Kumar PL, Sankung SB, Togola A, Tamò M. (2016) First report of outbreaks of the fall armyworm Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), a new alien invasive pest in west and central Africa. PloS ONE, 11(10), e0165632. DOI: https://doi.org/10.1371/journal. pone.0165632 . Guo JF, Zhang MD, Gao ZP, Wang DJ, He KL. and Wang, Z. Y. (2021).Comparison of larval performance and oviposition preference of Spodoptera frugiperda among three host plants: Potential risks to potato and tobacco crops. Insect Sci 28: 602–610. https://doi.org/10.1111/1744-7917.12830 . Haenniger S, Goergen G, Akinbuluma MD, Kunert M, Heckel DG, Unbehend M (2020) Sexual communication of Spodoptera frugiperd a from West Africa: Adaptation of an invasive species and implications for pest management. Sci Rep 10:1–9. He L, Zhao S, Gao X, Wu K (2021) Ovipositional responses of Spodoptera frugiperda on host plants provide a basis for using Bt-transgenic maize as trap crop in China. J Integr Agric 20: 804–814. Iken JE, Amusa NA (2004) Maize research and production in Nigeria. Afr J Biotechnol 3: 302–307. Johnson SJ (1987) Migration and the life history strategy of the fall armyworm, Spodoptera frugiperda in the western hemisphere. Int J Trop Insect Sci 8: 543–549. doi: 10.1017/S1742758400022591 . Karim ANMS, Ahmed S, Akhi AH, Talukder MZA, Mujahidi TA (2018) Combining ability and heterosis study in maize ( Zea mays L.) hybrids at different environments in Bangladesh. Bangladesh J Agric Res 43: 125–134. Khallaf MA, Knaden M (2020) Insect Host Choice: Don’t Put All the Eggs in One Basket. Curr Biol 30: 1363–1365. https://doi.org/10.1016/j.cub.2020.09.034 . Karavina C, Mandumbu R, Mukaro R (2014) Evaluation of three-way maize ( Zea mays L.) hybrids for yield and resistance to maize streak virus and turcicum leaf blight diseases. J Anim Plant Sci 24: 216–220. Kasoma C, Shimelis H, Laing MD (2021) Fall armyworm invasion in Africa: implications for maize production and breeding. J Crop Improv 35: 111–146. Khan Z, Sharawi SE, Khan MS, Suleman, Xing LX, Haroon, Ali S, Ahmed N (2022) Prevalence of insect pests on maize crop in District Mansehra, Khyber Pakhtunkhwa, Pakistan. Braz J Biol 84:e259217. doi: 10.1590/1519-6984.259217 . Kumar K, Gambhir G, Dass A, Tripathi AK, Singh A, Jha AK, Yadava P, Choudhary M, Rakshit S (2020) Genetically modified crops: Current status and future prospects. Planta 251:91. doi: 10.1007/s00425-020-03372-8 . Kumela T, Simiyu J, Sisay B, Likhayo P, Mendesil E, Gohole L, Tefera T. (2019) Farmers’ knowledge, perceptions, and management practices of the new invasive pest, fall armyworm ( Spodoptera frugiperda ) in Ethiopia and Kenya. Int J Pest Manage 65: 1–9. Kutka F (2011) Open-Pollinated vs. Hybrid Maize Cultivars. Sustainability 3: 1531–1554. Lee J, Chin JH, Ahn SN, Koh HJ (2015) Brief history and perspectives on plant breeding. In: Koh KH, Kwon SY, Thomson M (ed) Current Technologies in Plant Molecular Breeding: A Guide Book of Plant Molecular Breeding for Researchers, Springer, Dordrecht, pp. 1–14. Lenth R (2023) emmeans: Estimated Marginal Means, aka Least-Squares Means_. R package version 1.8.5, . Liebman M, Davis AS (2009) Managing weeds in organic farming systems: An ecological approach. In: Francis C (ed) Organic Farming: The Ecological System. Agronomy Monograph, Madison, WI, USA, pp 173–195. MacRobert JF, Setimela PS, Gethi J, Worku M (2014) Maize Hybrid Seed Production Manual. Mexico, D.F. CIMMYT. McCullagh P, Nelder JA (1989) Generalized Linear Models. London: Chapman and Hall. http://dx.doi.org/10.1007/978-1-4899-3242-6 . Meagher RL, Nagoshi RN, Stuhl CJ (2011) Oviposition Choice of Two Fall Armyworm (Lepidoptera: Noctuidae) Host Strains. J Insect Behav 24: 337–347. Montezano DG, Specht A, Sosa-Gómez DR, Roque-Specht VF, Sousa-Silva JC, Paula-Moraes SD, Hunt TE (2018) Host plants of Spodoptera frugiperda (Lepidoptera: Noctuidae) in the Americas. Afr Entomol 26: 286–300. Mutyambai DM, Niassy S, Calatayud PA, Subramanian S (2022) Agronomic Factors Influencing Fall Armyworm ( Spodoptera frugiperda ) Infestation and Damage and Its Co-occurrence with Stemborers in Maize Cropping Systems in Kenya. Insects 13: 266. https://doi.org/10.3390/insects13030266 . Nagoshi RN, Meagher RL, Hay-Roe M (2012) Inferring the annual migration patterns of fall armyworm (Lepidoptera:Noctuidae) in the United States from mitochondrial haplotypes. Ecol Evol 2: 1458–1467. Odeyemi OO, Lawal BO, Owolade OF, Olasoji JO, Egbetokun OA, Oloyede- Kamiyo QO, Omodele T, Anjorin FB (2020) Fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae): threat to maize production in Nigeria. Nig Agric J 51: 101–108. Ogunfunmilayo AO, Kazeem SA, Idoko JE, Adebayo RA, Fayemi EY, Adedibu OB, Ofuya TI (2021) Occurrence of natural enemies of fall armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae) in Nigeria. Plos one 16: 254–328. Ojumoola AO, Omoloye AA (2023) Agroecological zones influence maize infestation and damage severity by the fall armyworm (Spodoptera frugiperd a (J. E. Smith)) in southwestern Nigeria Acta Agric 119: 1–9. Olaniyan AB (2015) Maize: Panacea for hunger in Nigeria. Afr J Plant Sci 3: 155–174. Piepho HP (2004) An algorithm for a letter-based representation of all pairwise comparisons, J Comput Graph Stat 13: 456–466. Piesik D, Rochat D, Delaney KJ, Marion-Poll F (2012). Orientation of European corn borer first instar larvae to synthetic green leaf volatiles. J Applied Èntomol 137: 234–240. Pogue MG (2002) A world revision of the genus Spodoptera guenée (Lepidoptera: Noctuidae). Mem Am Entomol Soc 43:1–202. Prowell DP, McMichael M, Silvain JF (2004) Multilocus genetic analysis of host use, introgression, and speciation in host strains of fall armyworm (Lepidoptera: Noctuidae). Ann Entomol Soc Am 97:1034–1044. Pӧykkӧ H (2006) Females and larvae of a geometrid moth, Cleorodes lichenaria , prefer a lichen host that assures shortest larval period. Environ Entomol 35:1669–1675. R Core Team (2022) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/ . Refsnider JM, Janzen FJ (2010). Putting eggs in one basket: Ecological and evolutionary hypotheses for variation in oviposition-site choice. Annu Rev Ecol Evol Syst 41: 39–57. Rojas JC, Kolomiets MV, Bernal JS (2018) Nonsensical choices? Fall armyworm moths choose seemingly best or worst hosts for their larvae, but neonate larvae make their own choices. PloS one 13(5): e0197628. Rojas JC, Virgen A and Cruz-Lopez L (2003) Chemical and tactile cues influencing oviposition of a generalist moth, Spodoptera frugiperda (Lepidoptera: Noctuidae). Environ Entomol 32: 1386–1392. Russianzi W, Anwar R, Triwidodo H (2021) Biostatistics of fall armyworm Spodoptera frugiperda in maize plants in Bogor, West Java, Indonesia. Biodiversitas 22: 3463–3469. Rwomushana I, Bateman M, Beale T, Beseh P, Cameron K., Chiluba M, Tambo J (2018) Fall armyworm: impacts and implications for Africa. Wallingford, UK Saeed S, Sayyed AH, Ahmad I (2010) Effect of host plants on life-history traits of Spodoptera exigua (Lepidoptera: Noctuidae). J Pest Sci 83: 165–172. Sakamoto Y, Ishiguro M, Kitagawa G (1986) Akaike Information Criterion Statistics . D. Reidel Publishing Company. Singh GM, Xu J, Schaefer D, Day R, Wang Z, Zhang F (2021) Maize Diversity for Fall Armyworm Resistance in a Warming World. Crop Sci 62: 1–19. Sisay B, Sevgan S, Weldon CW, Krüger K, Torto B, Tamiru A. (2023) Responses of the fall armyworm ( Spodoptera frugiperda ) to different host plants: Implications for its management strategy. Pest Manage Sci 79: 845–856. Sobhy IS, Tamiru A, Chiriboga MX, Nyagol D, Cheruiyot D, Chidawanyika F, Subramanian S, Midega CAO, Bruce TJA, Khan ZR (2022) Bioactive Volatiles from Push-Pull Companion Crops Repel Fall Armyworm and Attract Its Parasitoids. Front Ecol Evol 10:883020. doi: 10.3389/fevo.2022.883020 . Sokame BM, Subramanian S, Kilalo DC, Juma G, Calatayud PA (2020) Larval dispersal of the invasive fall armyworm, Spodoptera frugiperda , the exotic stemborer Chilo partellus , and indigenous maize stemborers in Africa. Entomol Exp Appl, 168: 322–331. SoteloCardona P, Chuang WP, Lin MY, Chiang MY, Ramasamy S (2021) Oviposition preference not necessarily predicts offspring performance in the fall armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae) on vegetable crops. Sci Rep 11:15885; https://doi.org/10.1038/s41598-021-95399-4 . Sparks AN (1979) A review of the biology of the fall armyworm. Fla Entomol 62: 82e87. 10.2307/3494083 Szendrei Z, Rodriguez-Saona C (2010) A meta-analysis of insect pest behavioral manipulation with plant volatiles. Entomol Exp Appl 134: 201–210. Tambo JA, Kansiime MK, Mugambi I, Agboyi LK, Beseh PK, Day R (2023) Economic impacts and management of fall armyworm ( Spodoptera frugiperda ) in smallholder agriculture: a panel data analysis for Ghana. CABI Agric Biosci 4:38 https://doi.org/10.1186/s43170-023-00181-3 . Tiwari S (2022) Host plant preferences by fall armyworm Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae) on the range of potential host plant species. J Agric For 5: 25–33. Togola A, Meseka S, Menkir A, Badu-Apraku B, Boukar O, Tamò M, Djouaka R (2018) Measurement of Pesticide Residues from Chemical Control of the invasive Spodoptera frugiperda (Lepidoptera: Noctuidae) in a Maize Experimental Field in Mokwa, Nigeria. Int J Environ Res Public Health 15: 849–860. United States Department of Agriculture (USDA) (2020). World Agricultural Production: Global Market Analysis, International Production Assessment Division, Washington, DC https://apps.fas.usda.gov/psdonline/circulars/production.pdf . Volp TV, Zalucki MP, Furlong MJ (2022). What Defines a Host? Oviposition Behavior and Larval Performance of Spodoptera frugiperda (Lepidoptera: Noctuidae) on Five Putative Host Plants, J Econ Entomol 115: 1744–1751. Wang L, Yang A, He C, Qu M, Zhang J (2008). Creation of new maizegermplasm using alien introgression from Zea mays ssp. mexicana. Euphytica 164: 789–801. Yactayo-Chang JP, Mendoza J, Willams SD, Rering CC, Beck JJ, Block AK (2021). Zea mays Volatiles that Influence Oviposition and Feeding Behaviors of Spodoptera frugiperda. J Chem Ecol 47: 799–809. Yang X, Wyckhuys KA, Jia X, Nie F, Wu K. (2021). Fall armyworm invasion heightens pesticide expenditure among chinese smallholder farmers. J Environ Manage, 282, 111949. Yang G, Espelie KE, Wiseman BR, Ishenhour DJ (1993a) Effects of corn foliar cuticular lipids on the movement of fall armyworm (Lepidoptera: Noctuidae) neonate larvae. Fla Entomol 76: 302–316 Yang G, Wiseman BR, Ishenhour DJ, Espelie KE (1993b) Chemical and ultrastructural analysis of corn cuticular lipidsand their effects on fall armyworm larva. J Chem Ecol 9: 2055–2074. Yang J, Ma C, Jia R, Zhang H, Zhao Y, Yue H, Li H, Jiang X (2023) Different responses of two maize cultivars to Spodoptera frugiperda (Lepidoptera: Noctuidae) larvae infestation provide insights into their differences in resistance. Front Plant Sci 14:1065891. doi: 10.3389/fpls.2023.1065891 Zhang QY, Zhang YL, Quandahor, P., Gou, Y.P., Li, C.C., Zhang, K.X., Liu, C.Z. (2023). Oviposition Preference and Age-Stage, Two-Sex Life Table Analysis of Spodoptera frugiperda (Lepidoptera: Noctuidae) on different Maize Varieties. Insects 14: 413. https://doi.org/10.3390/insects14050413 . 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4601270","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":325665696,"identity":"ab295aa3-8cb7-4266-8a32-43114efacb99","order_by":0,"name":"Mobolade D. Akinbuluma","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2UlEQVRIiWNgGAWjYHACAwaGAxJAkrkByLGBiD0gTgsjSEsaRCyBsBYGmJbDhLXoNjBvYPhxxiLPXCKxTepGzfnEDccPsD3Ap8XsAFsBY88NiWLLGYlt0jnHbiduOJPAboBfC48BA88HicQNN0Ba2G4nzmxIYJMgpIXxD1zLv3OJM/sfENbCzHMDqiW37UBivwQhWw6zFRyWOSORuLPnYbN1bl+ycb/Ewzb8Wo43b3z45lhd4nb25IO3c77ZybbxJx+T+IBHCwMzOFZQADiCRsEoGAWjYBRQAgDDwVLHxGW8WAAAAABJRU5ErkJggg==","orcid":"","institution":"University of Ibadan","correspondingAuthor":true,"prefix":"","firstName":"Mobolade","middleName":"D.","lastName":"Akinbuluma","suffix":""},{"id":325665697,"identity":"e5de676e-228f-4ab4-9bac-46d26b2c03c0","order_by":1,"name":"Olubisi O. Bamifewe","email":"","orcid":"","institution":"University of Ibadan","correspondingAuthor":false,"prefix":"","firstName":"Olubisi","middleName":"O.","lastName":"Bamifewe","suffix":""},{"id":325665698,"identity":"73d3403e-642f-4060-80a4-be047c302d43","order_by":2,"name":"Olajumoke Y. Alabi","email":"","orcid":"","institution":"University of Ibadan","correspondingAuthor":false,"prefix":"","firstName":"Olajumoke","middleName":"Y.","lastName":"Alabi","suffix":""},{"id":325665699,"identity":"0d1bdaad-463b-4ab5-a5f6-3d8302206d73","order_by":3,"name":"Modupe I. J. Timothy","email":"","orcid":"","institution":"University of Ibadan","correspondingAuthor":false,"prefix":"","firstName":"Modupe","middleName":"I. J.","lastName":"Timothy","suffix":""},{"id":325665700,"identity":"08177449-44a3-4e88-a706-b127bfd0eb42","order_by":4,"name":"Peter Roessingh","email":"","orcid":"","institution":"University of Amsterdam","correspondingAuthor":false,"prefix":"","firstName":"Peter","middleName":"","lastName":"Roessingh","suffix":""},{"id":325665701,"identity":"e80aa246-5ef5-47c9-8e3c-538cafad97b2","order_by":5,"name":"Astrid T. Groot","email":"","orcid":"","institution":"University of Amsterdam","correspondingAuthor":false,"prefix":"","firstName":"Astrid","middleName":"T.","lastName":"Groot","suffix":""}],"badges":[],"createdAt":"2024-06-18 16:21:05","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4601270/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4601270/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":60171931,"identity":"c3d3e9ee-bfa5-4a3d-8941-bd03b7b8a6b0","added_by":"auto","created_at":"2024-07-12 15:14:08","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":78441,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of egg masses (± SEM, n = 4) of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e female. Line plots show number of egg masses on \u003cem\u003eDEKAIB, 30Y87, P3966W \u003c/em\u003e(hybrid maize varieties)\u003cstrong\u003e, LMFP, SWAN 1\u003c/strong\u003e and \u003cstrong\u003eV9928\u003c/strong\u003e (open pollinated varieties) over 72 h period on maize varieties in (A) multiple choice experiment and (B) no-choice experiments. ⃰ = significant difference between mean egg masses over 72 h of oviposition; ns = no significant differences between egg masses between the three time points (24, 48 and 72 h) of oviposition. Different letters under the error bars in a and b show that mean egg numbers between varieties are significantly different after 24h in multiple choice and no-choice experiments, respectively. C) Cumulative number of egg masses after 72 h of oviposition in multiple choice experiment and (D) cumulative number of egg masses in no-choice experiments. Boxes represent the lower and upper quartiles of egg masses on each maize, whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in c and d indicate significant differences among maize varieties (p \u0026lt; 0.05)\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4601270/v1/e5968e3ade82866475b74aa9.png"},{"id":60171925,"identity":"2a2c1e21-16a1-4852-a80d-9840f078ec01","added_by":"auto","created_at":"2024-07-12 15:14:08","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":83024,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of egg masses (± SEM, \u003cem\u003en = 4\u003c/em\u003e) of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e female laid on two types of maize varieties. Line plots show the number of egg masses on hybrid and OP maize varieties over a 72 h period in (A) multiple choice experiment and (B) no-choice experiment. Different letters on line show significant difference between mean egg masses laid over time. ⃰ in between the error bars in A and B show that mean egg numbers are significantly different after 24 h between hybrid and OP maize varieties in multiple choice (A) and no-choice experiment (B), respectively. (C) Mean cumulative number of egg masses over 72 h oviposition period in the multiple choice experiment and (D) in the no-choice experiment. Boxes represent the lower and upper quartiles of egg masses on each maize type; whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in C and D indicate significant differences between hybrid and OP varieties (p \u0026lt; 0.05)\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-4601270/v1/7560bb8e3fea3ff3903e405a.png"},{"id":60173573,"identity":"382c9a73-fc1b-4a61-b4d4-efaa5ee78709","added_by":"auto","created_at":"2024-07-12 15:22:08","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":19636,"visible":true,"origin":"","legend":"\u003cp\u003eOrientation responses of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003elarvae to odors from intact plants and clean air. Bar charts show the preferences of \u003cem\u003eS. frugiperda\u003c/em\u003e larvae to odors from intact four maize varieties and clean air presented in a Y-tube olfactometer. Hybrid maize varieties (\u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e) are in italics, Open pollinated maize varieties (\u003cstrong\u003eV9928, LMFP\u003c/strong\u003e) are in bold. Significant means were separated using Chi-square (χ2) test (p \u0026lt; 0.05, n=12), ns= not significant.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-4601270/v1/9debccdd76e05394650dfe37.png"},{"id":60174276,"identity":"8ed123c6-8e15-41d8-8397-a076e7fadc20","added_by":"auto","created_at":"2024-07-12 15:30:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":818341,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4601270/v1/63ecb92c-a310-4157-b898-678907cbf1ec.pdf"},{"id":60171928,"identity":"62f136a5-b07f-4667-98dd-d9a27be85ac7","added_by":"auto","created_at":"2024-07-12 15:14:08","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":57563,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryInformation.docx","url":"https://assets-eu.researchsquare.com/files/rs-4601270/v1/5e4ddb02aa1a443f6cfb0083.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Oviposition behavior and larval attraction of the fall armyworm Spodoptera frugiperda to different maize plant varieties","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe use of improved crop varieties is one of the key practices towards an efficient integrated pest management programme against herbivorous pests and pathogens (Acquaah \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Liebman and Davis \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). One focus of plant breeding programmes is generally to obtain hybrids that are able to withstand extreme conditions of cold or drought or that have resistance to pathogenic organisms or insect pests (Kutka \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Lee et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Kumar et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Enders and Begcy \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMaize (\u003cem\u003eZea mays\u003c/em\u003e L.) is a versatile multi-purpose crop, used as a feed globally and important food crop, especially in sub-Saharan Africa and Latin America (Erenstein et al. 2022). It is also one of the most important cereal crops in Nigeria grown for human consumption, animal feed and several industrial uses (Iken and Amusa \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Abdulraham and Kolawole \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Odeyemi et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Nigeria is the largest producer of maize in Africa with a production of 10.2\u0026nbsp;million metric tons on 4.8\u0026nbsp;million hectares of land, representing 0.42% of the total global production (FAO, 2018; Kamara et al. 2020). In maize, open-pollinated (OP) varieties are traditionally used varieties that inter-pollinate freely during seed production, resulting in heterogeneous varieties.\u003c/p\u003e \u003cp\u003eThe OP varieties have broad genetic bases selected by the environment and farmers over many generations, which help to maintain moderate stress resistance and yield characteristics (D\u0026aacute;vila-Flores et al. 2013; Lima et al. 2022). Thus, OP varieties are valuable genetic resources of breeding programmes (Costa et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Costa et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Dos Santos et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sing et al. 2021; Lima et al. 2022). In contrast to OP varieties, hybrid maize varieties are cultivated varieties that result from the fertilization of one maize plant by another genetically unrelated plant through controlled cross-pollination (FAO \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The quality of hybrid maize depends greatly on methods of field production, both in adherence to quality assurance standards and implementation of appropriate agronomic management (Bedő and Barnab\u0026aacute;s \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; MacRobert et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Karim et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Maize hybrid seeds provide farmers with varieties that have improved specific traits, such as high yield potential and unique trait combinations to counter diseases and adverse growing conditions (MacRobert et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), for example increased resistance to insect attack (Barry et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; de Lange et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and pathogens (Wang et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J.E. Smith) is a major phytophagous pest of agricultural crops endemic to tropical and subtropical regions of the Americas (Sparks, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Johnson et al. 1987; Nagoshi et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). It is a migrant pest with a wide host range causing great economic loss whenever present. The pest can feed on over 350 host plants belonging to 76 families, with the Poaceae being the most preferred host (Montezano et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Since \u003cem\u003eS. frugiperda\u003c/em\u003e invaded Africa (Georgen et al. 2016), it was reported in more than 45 African countries, affecting maize production (Kasoma et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Edosa et al. 2021) with over 50% of maize field losses (De Groote et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Abro et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In Nigeria, \u003cem\u003eS. frugiperda\u003c/em\u003e has established itself in three agroecological zones, causing extensive damage to maize farms, particularly in the humid forest agroecological zone in southwestern Nigeria (Ojumoola and Omoloye \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), causing between 50\u0026ndash;80% damage in many maize fields (Odeyemi et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne potential approach to \u003cem\u003eS. frugiperda\u003c/em\u003e management is to explore the role of maize varieties in influencing the attraction and acceptance behaviour of \u003cem\u003eS. frugiperda\u003c/em\u003e. Host plants play an important role in the chemical ecology and behaviour of \u003cem\u003eS. frugiperda\u003c/em\u003e, because the insects respond to chemical cues for oviposition and larval feeding (Guo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Sotelo-Cardona et al. 2021; Zhang et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In general, different maize varieties, whether hybrid or OP, vary in their biochemical composition, which make them more or less suitable to \u003cem\u003eS. frugiperda\u003c/em\u003e (Yang et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). For example, some maize varieties produce higher levels of secondary metabolites such as terpenoids, which have been shown to have insecticidal properties, while other varieties may produce compounds that are attractive to \u003cem\u003eS. frugiperda\u003c/em\u003e (Firake et al. 2020). In addition, physical characteristics of host plants, such as trichomes, wax amount, thickness and toughness of leaves and secondary toxic metabolites influence host-plant selection behavior (Gatehouse \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Differential preference of \u003cem\u003eS. frugiperda\u003c/em\u003e was reported in maize cultivars due to difference in cuticular lipids (Yang 1993a), presence of wax materials on the leaf surface (Yang 1993b) and anti-feedant and anti-repellent properties (Tiwari \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral studies have investigated the oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e to identify which host plant species are either resistant or susceptible to the insect pest (Ba et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Tiwari \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Sisay et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Only few studies evaluated oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e on different maize varieties (He et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The use of maize varieties that are less preferred by \u003cem\u003eS. frugiperda\u003c/em\u003e could reduce the level of infestation in the field. Whether Nigerian \u003cem\u003eS. frugiperda\u003c/em\u003e females show any oviposition preference towards specific hybrid or OP maize varieties is largely unknown. In addition, little is known about the larval attraction of \u003cem\u003eS. frugiperda\u003c/em\u003e to different maize plant varieties, particularly in Nigeria (Odeyemi et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo evaluate whether \u003cem\u003eS. frugiperda\u003c/em\u003e show differential attraction and oviposition behaviours towards different maize varieties, we investigated the oviposition preference of the Nigerian \u003cem\u003eS. frugiperda\u003c/em\u003e population on six maize varieties, three of which were hybrid and three were OP varieties. To determine whether the number of egg masses laid by \u003cem\u003eS. frugiperda\u003c/em\u003e female increased or decreased over the oviposition period, we checked for variability in the timing of egg-laying on the maize varieties, by conducting oviposition assay over three days in both multiple choice and no-choice experiments. Finally, we determined whether the \u003cem\u003eS. frugiperda\u003c/em\u003e larvae were attracted differentially towards the odors from the different maize plant varieties.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy location\u003c/h2\u003e \u003cp\u003eThe experiment was conducted in the Entomology Research Laboratory and Insect Chemical Ecology Laboratory, Department of Crop Protection and Environmental Biology, University of Ibadan, Nigeria, under ambient conditions of 27\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C and 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eSource of maize seeds and planting\u003c/h2\u003e \u003cp\u003eThree hybrid maize varieties (referred to in italics in the remainder of this manuscript), \u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e, and three OP varieties (referred to in bold), \u003cb\u003eLMFP, SWAN1\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e, were used for experiments. \u003cem\u003eDEKALB\u003c/em\u003e was obtained from International Institute of Tropical Agriculture; \u003cb\u003eV9928, LMFP\u003c/b\u003e and \u003cb\u003eSWAN1\u003c/b\u003e were obtained from Institute of Agriculture Research and Training, Moor Plantation, Apata Ibadan, while \u003cem\u003e30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e were obtained from Corteva Agriscience. These varieties were selected because they are cultivated by Nigerian farmers, but so far were never tested for attraction and oviposition by \u003cem\u003eS. frugiperda\u003c/em\u003e. Two seeds from each variety were sown in a 10 kg pot filled with heat-sterilized loamy soil. The potted plants were grown and maintained in a green house at 26\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and a photoperiod of 14:10 (L:D) and each variety was planted in six pots. To have a sufficient number of plants for the experiments, the plants were sown every week. Seven days after each sowing, maize plants in each pot were thinned to one. For the experiments, 14-day old potted maize plants were used.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eInsect collection and rearing\u003c/h2\u003e \u003cp\u003e \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e larvae were collected from a naturally infested maize farms at Sasha-Ajibode (latitude 7\u003csup\u003eo\u003c/sup\u003e 28\u003csup\u003e1\u003c/sup\u003e 37.70688 E, 3\u003csup\u003e0\u003c/sup\u003e 54\u003csup\u003e1\u003c/sup\u003e N) and Elekuru-Akinyele areas (7\u003csup\u003e0\u003c/sup\u003e 36\u003csup\u003e1\u003c/sup\u003e25.54092,3\u003csup\u003e0\u003c/sup\u003e 49\u003csup\u003e1\u003c/sup\u003e N) of Ibadan, Southwestern Nigeria and reared at the Entomology Research Laboratory, University of Ibadan, Nigeria. The larvae were separated into a transparent sauce cup (EEZEE, Nigeria; 4 cm \u0026times;3 cm; 40 mL) with one larva per cup, which were fed daily with fresh maize leaves of the SWAN 1 variety and covered with lid until pupation. The pupae were placed into clean vials lined with moist tissue paper until adult emergence. The adult insects were kept in the same vials and fed with 10% sugar solution. For mating to occur, 2\u0026ndash;3 day old virgin adults were paired in mating cups (AVT Plastics, 500 mL, one pair/cup) with a sauce cup (4 cm \u0026times;3 cm; 40 mL) filled with cotton wool soaked in a 10% sugar solution and covered with muslin cloth. Eggs were collected daily and newly emerged larvae were individually placed in cups and fed with fresh maize leaves of the SWAN 1 variety (approx. 1 g/cup) for 7\u0026ndash;12 days. For the experiments, 2\u0026ndash;4 day-old mated adult females and the 7\u0026ndash;12 day old larvae of the new generations were used.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eOviposition assays\u003c/h2\u003e \u003cp\u003eOviposition preference and performance of \u003cem\u003eS. frugiperda\u003c/em\u003e females was determined in multiple-choice and no-choice experiments inside cages enclosed in the same green house as described above, using intact plants from the six maize varieties. In the multiple choice test, six pots were properly labeled and placed in a mesh cage (40 \u0026times; 35 \u0026times; 50 cm), whereby each pot contained one 14 day-old plant of one maize variety. The potted plants were arranged inside the cage in a completely randomized design (CRD). One 2\u0026ndash;4 day-old mated female was released into the cage for 72 hours. The multiple choice test was conducted in four replicates. Each replicate contained different batch of the six plants (of the same age) and was set up separately at one week interval. The number of egg masses on the maize plants inside the cage was counted and recorded every 24 h for a period of 72 h.\u003c/p\u003e \u003cp\u003eIn the no-choice experiment, we placed one pot containing one 14 day-old plant of each variety within similar mesh cages (40 \u0026times; 35 \u0026times; 50 cm) together with one 2\u0026ndash;4 day-old mated female. The no-choice experiment thus had six treatments with 4 replications for each variety. The number of egg masses on the maize plants was counted and recorded every 24 h for a period of 72 h. Egg masses were carefully counted and recorded from outside the cage such that female insects were not disturbed. All data were collected between 13:00\u0026ndash;16:00 daily and the bioassays were conducted between March and June 2022.\u003c/p\u003e \u003cp\u003e \u003cb\u003eResponses of\u003c/b\u003e \u003cb\u003eSpodoptera frugiperda\u003c/b\u003e \u003cb\u003elarvae to intact plants\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo determine the orientation responses of \u003cem\u003eS. frugiperda\u003c/em\u003e larvae to intact maize plants, we chose to test the two most preferred varieties and two least preferred varieties based on the oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e females from the experiment above. The experiment was conducted in a Y-tube olfactometer, as described in Akinbuluma and Chinaka (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) with some modifications, and summarized here. The experimental arena consisted of a horizontal pyrex glass Y-tube (10 mm i.d; stem 85 mm; arms 75 mm at a 60\u0026deg; angle to the stem). Air from a field pump was passed through activated charcoal and humidified with double distilled water. The airflow was split into two halves. One half was passed through a glass chamber with a tightly sealed lid, enclosing a potted plant (test) and into one arm of the olfactometer, while the other half of the airflow was passed through an empty glass chamber (control) at the same regulated flow rate of 60 mL/min. A vacuum line was connected to a mini pump powered by a rechargeable battery pulled air through the two arms of the Y-tube. Fourth instar \u003cem\u003eS. frugiperda\u003c/em\u003e larvae were placed in the bioassay room for at least 12 h before the experiment to acclimatize them to the room conditions. The larvae were also starved for 3 hours before the experiment by removing the larvae from their feed container to a small empty rearing cup (1 larva/cup). Twelve larvae were individually assayed in random order on the plant varieties such that 1 larva was assessed 4 times on the four maize varieties. Larval choice was recorded after spending 3 mins in any of the arms of the Y-tube. Insects that failed to choose an arm within 10 mins were recorded as non-responders and were not included in analysis.\u003c/p\u003e \u003cp\u003eAfter testing on each maize variety, the olfactometer set-up, glass chambers and connections were wiped with 70% ethanol and dried to avoid odor contamination between consecutive bioassays and the odor source positions were also exchanged. After an interval of 15 mins, the next larva was tested. Bioassays were always conducted between 13:00 and 18:00 hours. This study was conducted between May and July 2022.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eStatistics and Data Analysis\u003c/h2\u003e \u003cp\u003eThe multiple-choice and no-choice oviposition experiments were analyzed separately. The number of egg masses was modeled with generalized linear models (GLM, McCullagh and Nelder, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1989\u003c/span\u003e) using Poisson regression. Both models contained maize variety and duration of oviposition as well as their interaction as explanatory variables. In addition, the position of the egg masses in the multiple choice experiment was determined using a separate model, containing position, hours and their interaction. Finally we investigated if there were differences between the Hybrid varieties (\u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e) and OP varieties (\u003cb\u003eLMFP, SWAN1\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e) with a model that contained the varietal type and hour, as well as their interaction as explanatory variables (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Posthoc comparisons and testing the trends of egg laying over time were done on marginal means (Lenth \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and the result displayed with the compact letter display (Piepho, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), with p\u0026thinsp;=\u0026thinsp;0.05 as significance level and Tukey adjustment for multiple testing. The olfactometer experiment data were analyzed using chi-square tests. All statistical analyses were performed in R (R Core Team \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eOverview of the explanatory factors in models used. In all models, the response was the number of egg masses laid by female \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003cp\u003e(Explanatory variables)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eN\u003c/p\u003e \u003cp\u003e(cases)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDf\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChoice test\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVariety * hour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e64\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePosition of egg masses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePosition * hour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNo-choice test\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVariety * hour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e64\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMaize-type test\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eType * hour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e54\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003e \u003cb\u003eOviposition preference of adult\u003c/b\u003e \u003cb\u003eS. frugiperda\u003c/b\u003e \u003cb\u003efemale\u003c/b\u003e\u003c/p\u003e \u003cp\u003eWhen we checked the number of egg masses laid by \u003cem\u003eS. frugiperda\u003c/em\u003e female on the different maize varieties in the multiple choice experiment, we found significant difference between the maize varieties (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05, n\u0026thinsp;=\u0026thinsp;72, df\u0026thinsp;=\u0026thinsp;64), but no significant interaction between hours of oviposition and maize variety.\u003c/p\u003e \u003cp\u003eSignificantly fewer egg masses were laid over the 72-hour period on \u003cem\u003eP3966W\u003c/em\u003e and \u003cb\u003eLMFP\u003c/b\u003e but not on \u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e, \u003cb\u003eSWAN 1\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). However, after 24 h of oviposition, the number of egg masses found on \u003cem\u003eDEKAIB\u003c/em\u003e was significantly lower than those of \u003cem\u003eP3966W\u003c/em\u003e, \u003cb\u003eLMFP\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e varieties (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). Similarly, in the no-choice experiment, we found significant difference between the maize varieties but no interaction with hours of oviposition (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05, n\u0026thinsp;=\u0026thinsp;72, df\u0026thinsp;=\u0026thinsp;64). Across 72 h oviposition period, we observed significantly fewer egg masses (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) of \u003cem\u003eS. frugiperda\u003c/em\u003e on maize varieties, \u003cb\u003eLMFP\u003c/b\u003e and \u003cb\u003eSWAN 1\u003c/b\u003e, but not on \u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB). However, there was no significant difference in the number of egg masses laid across all maize varieties after 24 h of oviposition (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003eWhen we combined the egg masses over time for each maize variety in the multiple choice experiment (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB), we found that \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e had significantly lower egg mass numbers than varieties, \u003cb\u003eLMFP, SWAN 1\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). However, similar numbers of egg mass was found on \u003cem\u003e30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e (p\u0026thinsp;\u0026gt;\u0026thinsp;0.05). Similarly, in the no-choice experiment, we found significantly lower egg mass number on \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e than on \u003cb\u003eLMFP\u003c/b\u003e maize varieties (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eD).\u003c/p\u003e \u003cp\u003eWhen combining all three hybrid varieties and the OP varieties, egg mass counts only significantly differed between 24 h and 72 h in the multiple choice (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA), with no significant difference in the number of eggs laid over 72 h in the no-choice experiment (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA and B). In the OP varieties, we found significantly lower egg masses on the hybrid varieties after 72 h of oviposition than after 24 h both in the multiple choice and the no-choice test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). After 24 h of oviposition, significantly fewer egg masses were laid by \u003cem\u003eS. frugiperda\u003c/em\u003e female on hybrid maize than on OP varieties in both experiments (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA and B). When the cumulative egg mass counts were compared, egg masses on the hybrid varieties were significantly lower than those on the OP varieties in both multiple choice and no-choice experiments (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eC, D).\u003c/p\u003e \u003cp\u003e \u003cb\u003eResponse of\u003c/b\u003e \u003cb\u003eSpodoptera frugiperda\u003c/b\u003e \u003cb\u003elarvae to intact maize varieties\u003c/b\u003e\u003c/p\u003e \u003cp\u003eOlfactometer experiment with intact plants of the four maize varieties showed that significantly more \u003cem\u003eS. frugiperda\u003c/em\u003e larvae (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;8.33; p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) chose LMFP maize variety than clean air (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Larvae did not show a preference to the other three tested maize varieties relative to clean air (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.33; p\u0026thinsp;\u0026gt;\u0026thinsp;0.05), 30Y87 (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.33; p\u0026thinsp;\u0026gt;\u0026thinsp;0.05) and DEKAIB (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.33; p\u0026thinsp;\u0026gt;\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e Mean number of egg masses (\u0026plusmn;\u0026thinsp;SEM, n\u0026thinsp;=\u0026thinsp;4) of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e female. Line plots show number of egg masses on \u003cem\u003eDEKAIB, 30Y87, P3966W\u003c/em\u003e (hybrid maize varieties), \u003cb\u003eLMFP, SWAN 1\u003c/b\u003e and \u003cb\u003eV9928\u003c/b\u003e (open pollinated varieties) over 72 h period on maize varieties in (A) multiple choice experiment and (B) no-choice experiments. * = significant difference between mean egg masses over 72 h of oviposition; ns\u0026thinsp;=\u0026thinsp;no significant differences between egg masses between the three time points (24, 48 and 72 h) of oviposition. Different letters under the error bars in a and b show that mean egg numbers between varieties are significantly different after 24h in multiple choice and no-choice experiments, respectively. C) Cumulative number of egg masses after 72 h of oviposition in multiple choice experiment and (D) cumulative number of egg masses in no-choice experiments. Boxes represent the lower and upper quartiles of egg masses on each maize, whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in c and d indicate significant differences among maize varieties (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e Mean number of egg masses (\u0026plusmn;\u0026thinsp;SEM, \u003cem\u003en\u0026thinsp;=\u0026thinsp;4\u003c/em\u003e) of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e female laid on two types of maize varieties. Line plots show the number of egg masses on hybrid and OP maize varieties over a 72 h period in (A) multiple choice experiment and (B) no-choice experiment. Different letters on line show significant difference between mean egg masses laid over time. * in between the error bars in A and B show that mean egg numbers are significantly different after 24 h between hybrid and OP maize varieties in multiple choice (A) and no-choice experiment (B), respectively. (C) Mean cumulative number of egg masses over 72 h oviposition period in the multiple choice experiment and (D) in the no-choice experiment. Boxes represent the lower and upper quartiles of egg masses on each maize type; whiskers indicate the minimum and maximum values, middle line represents the median. Different letters above the bars in C and D indicate significant differences between hybrid and OP varieties (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e Orientation responses of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e larvae to odors from intact plants and clean air. Bar charts show the preferences of \u003cem\u003eS. frugiperda\u003c/em\u003e larvae to odors from intact four maize varieties and clean air presented in a Y-tube olfactometer. Hybrid maize varieties (\u003cem\u003eDEKAIB, 30Y87\u003c/em\u003e) are in italics, Open pollinated maize varieties (\u003cb\u003eV9928, LMFP\u003c/b\u003e) are in bold. Significant means were separated using Chi-square (χ2) test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05, n\u0026thinsp;=\u0026thinsp;12), ns\u0026thinsp;=\u0026thinsp;not significant.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we investigated the oviposition and attraction of \u003cem\u003eS. frugiperda\u003c/em\u003e to different maize varieties, belonging to hybrid and OP varieties, and found i) variation in the number of egg masses laid on maize varieties at different time periods, ii) ovipositional preferences of \u003cem\u003eS. frugiperda\u003c/em\u003e females for \u003cb\u003eSWAN 1, V9928\u003c/b\u003e and \u003cb\u003eLMFP\u003c/b\u003e compared to \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e in both multiple-choice and no-choice experiments, iii) significant attraction responses of \u003cem\u003eS. frugiperda\u003c/em\u003e larvae towards \u003cb\u003eLMFP\u003c/b\u003e maize variety compared to clean air, while this was not the case for \u003cb\u003eV9928\u003c/b\u003e, \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e. Overall, our results show that DEKAIB and 30Y87 were less accepted and less attractive to \u003cem\u003eS. frugiperda\u003c/em\u003e than the other maize varieties.\u003c/p\u003e \u003cp\u003eOur results showed significant variation in the number of egg masses laid by \u003cem\u003eS. frugiperda\u003c/em\u003e females on the different maize varieties, both in the multiple choice and no-choice tests. The consistently significant lower number of egg masses on \u003cem\u003eDEKAIB\u003c/em\u003e suggests that the variety was the least acceptable for oviposition compared to all other tested varieties. Differential oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e female for some maize varieties has been shown by others as well (He et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and could be due to differences in plant nutrients in maize host plants, which is important for development of offspring (Tiwari \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor a proper timing of management of \u003cem\u003eS. frugiperda\u003c/em\u003e eggs on the maize host plants, it is important to know when egg masses are laid and how this changes over time. \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e females have been found to lay reduced numbers of egg batches over time (Russianzi et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Therefore, we counted the number of egg masses over a three-day period. Among the varieties tested, we found significant reductions in the number of egg masses laid over time. For example, in the no-choice test, significantly more egg batches were laid on the \u003cb\u003eLFMP\u003c/b\u003e and \u003cb\u003eSWAN1\u003c/b\u003e maize varieties after 24 h than after 72 h. Also, in the first 24 h, significantly less egg batches were laid on the hybrid maize varieties (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). We also showed that significantly less egg masses were found on the combined hybrid maize varieties \u003cem\u003e(DEKAIB, 30Y87\u003c/em\u003e and \u003cem\u003eP3966W\u003c/em\u003e) than on the OP varieties (\u003cb\u003eLMFP, V9928\u003c/b\u003e and \u003cb\u003eSWAN1\u003c/b\u003e). Even when considered separately, the least number of egg masses were laid on the hybrid varieties compared to the OP varieties (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Our results thus indicate that hybrid maize varieties are less favourable to \u003cem\u003eS. frugiperda\u003c/em\u003e for oviposition than OP varieties. An earlier report also showed that \u003cem\u003eS. frugiperda\u003c/em\u003e did not infest hybrid maize varieties as much as OP varieties in both the lowland and high-altitude land in Kenya (Mutyambai et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, Zhang et al. (\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) found significant oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e females for some hybrid maize varieties compared to OP varieties. Thus, differential oviposition cannot always be explained by maize pollination type. Nevertheless, to reduce infestation of maize by \u003cem\u003eS. frugiperda\u003c/em\u003e in Nigeria, our results suggest that farmers should use hybrid maize varieties instead of OP varieties for establishing maize fields.\u003c/p\u003e \u003cp\u003eWe would like to point out that in all our oviposition experiments, we found more \u003cem\u003eS. frugiperda\u003c/em\u003e egg masses on the cage surfaces than the maize varieties (Supplementary Fig. SI). This is a common phenomenon that other studies also found (Rojas et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Barros et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Meagher et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Sotelo-Cardona et al. 2021; Volp et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This may suggest that some maize plants volatiles have repellent effects for female insects, preventing them from ovipositing on the plants. Rojas et al. (\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and Barros et al. (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) reported that \u003cem\u003eS. frugiperda\u003c/em\u003e female oviposited more on corrugated surfaces rather than surfaces treated with some host plant extracts. Thus, our study corroborates the finding that \u003cem\u003eS. frugiperda\u003c/em\u003e females lay more eggs on oviposition cage walls away from nearby plants than on plants, which could indicate repellent effects of some plant-derived volatiles (Sisay et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe olfactometer bioassay with \u003cem\u003eS. frugiperda\u003c/em\u003e larvae showed only a larval preference towards \u003cb\u003eLMFP\u003c/b\u003e maize variety relative to clean air, and not to the other varieties. Possibly, this variety releases volatiles that are attractive for larvae. Although lepidopteran larvae possess limited mobility, they can still move between plants by way of so-called \u0026lsquo;ballooning\u0026rsquo; (e.g. Sokame et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and also orient towards or away from host plants (Liu et al. 2022). Background volatile compounds have been found to modify larval behavior in \u003cem\u003eS. frugiperda\u003c/em\u003e on some maize varieties (Yactayo-Chang et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Interestingly, \u003cem\u003eS. frugiperda\u003c/em\u003e larvae were not significantly attracted to the hybrid \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e, on which \u003cem\u003eS. frugiperda\u003c/em\u003e females oviposited least.\u003c/p\u003e \u003cp\u003eIn conclusion, our study showed that different maize varieties elicit differential oviposition and larval responses in \u003cem\u003eS. frugiperda.\u003c/em\u003e Possibly, variation in maize host plant volatiles may partly account for these different responses. Identification of plant volatiles that attract \u003cem\u003eS. frugiperda\u003c/em\u003e females could help to reduce the population. Since the use of plant varieties can affect the extent of infestation and damage by insect pest, varieties with less preference for oviposition by the insect will suffer less infestation and damage compared to those with full acceptance for oviposition by the insect. Our study suggests that planting \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e, which are hybrid maize varieties could reduce the level of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e infestation in Nigerian fields. Exploring these avenues further will help to develop integrated pest management (IPM) strategies of this pest.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eConflicts of interest/Competing interests\u003c/h2\u003e\n\u003cp\u003eThe authors declare that they have no conflict of interest/competing interests\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSupplementary Information\u003c/strong\u003e Supplementary information can be found on the online version of this article.\u003c/p\u003e\n\u003ch2\u003eAvailability of data and materials\u003c/h2\u003e\n\u003cp\u003eThe authors confirm that data supporting the findings of this study are available from the corresponding author on request.\u003c/p\u003e\n\u003ch2\u003eFunding:\u003c/h2\u003e\n\u003cp\u003eThis project was part of the funded project by a grant from the International Foundation for Science, co-sponsored by the Organisation for the Prohibition of Chemical Weapons (OPCW).\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eM.D.A planned and conceived the study and acquired funding. MDA, OOB and MIJT conducted the experiments. OYA contributed towards olfactometer setup. MDA and PR performed data analysis. MDA and OOB wrote the draft manuscript. MDA, PR and ATG reviewed the manuscript. All authors read and approved the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eWe are grateful to the International Foundation for Science, co-sponsored by the Organisation for the Prohibition of Chemical Weapons (OPCW), for funding this work. We thank Dr Felicia O. Aigbokhan of the Corteva Agrisciences, for facilitating the provision of some hybrid maize seeds for this research.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbdulraham AA, Kolawole OM (2006) Traditional preparation and uses of maize in Nigeria. Ethnobotany Leaflets 10: 291\u0026ndash;227.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAbro Z, Kimathi E, De Groote H, Tefera T, Sevgan S, Niassy S, Kassie M (2021) Socioeconomic and health impacts of fall armyworm in Ethiopia PLoS ONE, 16, e0257736.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAcquaah G (2004) Principles of Crop Production: Theory, Techniques, and Technology. Pearson Education, Prentice Hall: Upper Saddle River, NJ, USA.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAkinbuluma MD, Chinaka OP (2023) Efficacy of the parasitic wasp, \u003cem\u003eDinarmus basalis\u003c/em\u003e Rondani (Hymenoptera: Pteromalidae), in reducing infestations by the cowpea beetle, \u003cem\u003eCallosobruchus maculatus\u003c/em\u003e (L.) (Coleoptera: Chrysomelidae: Bruchinae). Egypt J Biol Pest Control 33: 1\u0026ndash;7.\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s41938-023-00692-1\u003c/span\u003e\u003cspan address=\"10.1186/s41938-023-00692-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBa TX, Zhang YH, Zhang Z, Guan DD, Li CC, Ji ZY, Yin XT, Zhang AH, Tang QB, Liu YH, Li XR, Zhou X (2020) The host preference and population life tables of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) fed on maize and wheat. Plant Prot 46:17\u0026ndash;23.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBarros EM, Torres JB, Ruberson JR, Oliveira MD (2010) Development of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e on different hosts and damage to reproductive structures in cotton. Entomol Exp Appl 137: 237\u0026ndash;245.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBarry D, Widstrom NW, Darrah LL, McMillian WW, Riley TJ, Scott GE, Lillehoj EB (1992) Maize ear damage by insects in relation to genotype and aflatoxin contamination in preharvest maize grain. J Econ Entomol 85: 2492\u0026ndash;2495.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBedő Z, Barnab\u0026aacute;s B (2013) 60 Years of Hungarian Hybrid Maize 1953\u0026ndash;2013, Pannonian Plant Biotechnology Association, Martonv\u0026aacute;s\u0026aacute;r, Hungary. pp. 135.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCarroll MJ, Schmelz EA, Meagher RL, Teal PE (2006) Attraction of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e larvae to volatiles from herbivore-damaged maize seedlings. J Chem Ecol 32: 1911\u0026ndash;1924.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChimweta M, Nyakudya IW, Jimu L., Bray Mashingaidze A (2020). Fall armyworm [\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J. E Smith)] damage in maize: management options for food-recession cropping smallholder farmers Int J Pest Manag 66:142\u0026ndash;154.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCosta EN, Fernandes MG, Medeiros PH, Evangelista BMD (2020) Resistance of Maize Landraces from Brazil to Fall Armyworm (Lepidoptera: Noctuidae) in the Winter and Summer Seasons. Bragantia 79: 377\u0026ndash;386.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCosta EN, Nogueira L, De Souza BHS, Ribeiro ZA, Louvandini H, Zukoff SN, J\u0026uacute;nior ALB (2018) Characterization of Antibiosis to \u003cem\u003eDiabrotica speciosa\u003c/em\u003e (Coleoptera: Chrysomelidae) in Brazilian Maize Landraces. J Econ Entomol 111: 454\u0026ndash;462.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCrabb R (1992) The Hybrid Corn-Makers. West Chicago Publishing Co. Chicago, IL, USA.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDay R, Abrahams P, Bateman M, Beale T, Clottey V, Cock M, Witt A (2017) Fall armyworm: impacts and implications for Africa. Outlooks on Pest Manage 28: 196\u0026ndash;201.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Groote H, Kimenju SC, Munyua B, Palmas S, Kassie M, Bruce A (2020). Spread and impact of fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e J. E. Smith) in maize production areas of Kenya. Agriculture, Ecosystem and Environment, 292, 106804.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ede Lange ES, Balmer D, Mauch-Mani B, Turlings TCJ (2014) Insect and pathogen attack and resistance in maize and its wild ancestors, the teosintes. New Phytol 204: 329\u0026ndash;341.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDespland E (2021) Selection Forces Driving Herding of Herbivorous Insect Larvae. Front Ecol Evol 9:760806. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3389/fevo.2021.760806\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2021.760806\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDos Santos LFC, Ruiz-S\u0026aacute;nchez E, Andueza-Noh RH, Garru\u0026ntilde;a-Hern\u0026aacute;ndez R, Latournerie-Moreno L, Mijangos-Cort\u0026eacute;s JO (2020) Leaf Damage by \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e J. E. Smith (Lepidoptera: Noctuidae) and Its Relation to Leaf Morphological Traits in Maize Landraces and Commercial Cultivars. J Plant Dis Protect 127: 103\u0026ndash;109.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDuvick DN, Smith JSC, Cooper M. (2004) Changes in performance, parentage, and genetic diversity of successful corn hybrids 1930\u0026ndash;2000. In: Smith CW, Betran J, Runge ECA (ed) Corn: Origin, History, Technology and Production, John Wiley and Sons, Hoboken, NJ, USA, pp 65\u0026ndash;97.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEdosa TT, Dinka TD (2021) Current and future potential distribution, risk and management of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e. J Innov Agric 8: 14\u0026ndash;23.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEnders L, Begcy K. (2021) Unconventional routes to developing insect-resistant crops. Mol Plant 14: 439\u0026ndash;1453.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eErenstein O, Jaleta M, Sonder K, Mottaleb K, Prasanna BM. (2002) Global maize production, consumption and trade: trends and R\u0026amp;D implications. Food Secur 14: 1295\u0026ndash;1319.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFAO (2016) Seed Security Assessment: Handout 0 - Session 3 (Seed systems). Pages 1\u0026ndash;3. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.fao.org/agriculture/crops/core-themes/theme/seeds-pgr/know_res/en/\u003c/span\u003e\u003cspan address=\"http://www.fao.org/agriculture/crops/core-themes/theme/seeds-pgr/know_res/en/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFirake DM, Behere GT (2020) Bioecological attributes and physiological indices of invasive fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J.E. Smith) infesting ginger (\u003cem\u003eZingiber officinale\u003c/em\u003e Roscoe) plants in India. Crop Prot 137: 105\u0026ndash;233.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGatehouse JA (2002) Plant resistance towards insect herbivores: A dynamic interaction. New Phytol 156:145\u0026ndash;169.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGoergen G, Kumar PL, Sankung SB, Togola A, Tam\u0026ograve; M. (2016) First report of outbreaks of the fall armyworm \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J.E. Smith) (Lepidoptera: Noctuidae), a new alien invasive pest in west and central Africa. PloS ONE, 11(10), e0165632. DOI: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal. pone.0165632\u003c/span\u003e\u003cspan address=\"10.1371/journal. pone.0165632\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGuo JF, Zhang MD, Gao ZP, Wang DJ, He KL. and Wang, Z. Y. (2021).Comparison of larval performance and oviposition preference of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e among three host plants: Potential risks to potato and tobacco crops. Insect Sci 28: 602\u0026ndash;610. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1744-7917.12830\u003c/span\u003e\u003cspan address=\"10.1111/1744-7917.12830\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHaenniger S, Goergen G, Akinbuluma MD, Kunert M, Heckel DG, Unbehend M (2020) Sexual communication of \u003cem\u003eSpodoptera frugiperd\u003c/em\u003ea from West Africa: Adaptation of an invasive species and implications for pest management. Sci Rep 10:1\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHe L, Zhao S, Gao X, Wu K (2021) Ovipositional responses of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e on host plants provide a basis for using Bt-transgenic maize as trap crop in China. J Integr Agric 20: 804\u0026ndash;814.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eIken JE, Amusa NA (2004) Maize research and production in Nigeria. Afr J Biotechnol 3: 302\u0026ndash;307.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJohnson SJ (1987) Migration and the life history strategy of the fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e in the western hemisphere. Int J Trop Insect Sci 8: 543\u0026ndash;549. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1017/S1742758400022591\u003c/span\u003e\u003cspan address=\"10.1017/S1742758400022591\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKarim ANMS, Ahmed S, Akhi AH, Talukder MZA, Mujahidi TA (2018) Combining ability and heterosis study in maize (\u003cem\u003eZea mays\u003c/em\u003e L.) hybrids at different environments in Bangladesh. Bangladesh J Agric Res 43: 125\u0026ndash;134.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKhallaf MA, Knaden M (2020) Insect Host Choice: Don\u0026rsquo;t Put All the Eggs in One Basket. Curr Biol 30: 1363\u0026ndash;1365. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.cub.2020.09.034\u003c/span\u003e\u003cspan address=\"10.1016/j.cub.2020.09.034\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKaravina C, Mandumbu R, Mukaro R (2014) Evaluation of three-way maize (\u003cem\u003eZea mays\u003c/em\u003e L.) hybrids for yield and resistance to maize streak virus and turcicum leaf blight diseases. J Anim Plant Sci 24: 216\u0026ndash;220.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKasoma C, Shimelis H, Laing MD (2021) Fall armyworm invasion in Africa: implications for maize production and breeding. J Crop Improv 35: 111\u0026ndash;146.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKhan Z, Sharawi SE, Khan MS, Suleman, Xing LX, Haroon, Ali S, Ahmed N (2022) Prevalence of insect pests on maize crop in District Mansehra, Khyber Pakhtunkhwa, Pakistan. Braz J Biol 84:e259217. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1590/1519-6984.259217\u003c/span\u003e\u003cspan address=\"10.1590/1519-6984.259217\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKumar K, Gambhir G, Dass A, Tripathi AK, Singh A, Jha AK, Yadava P, Choudhary M, Rakshit S (2020) Genetically modified crops: Current status and future prospects. Planta 251:91. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s00425-020-03372-8\u003c/span\u003e\u003cspan address=\"10.1007/s00425-020-03372-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKumela T, Simiyu J, Sisay B, Likhayo P, Mendesil E, Gohole L, Tefera T. (2019) Farmers\u0026rsquo; knowledge, perceptions, and management practices of the new invasive pest, fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e) in Ethiopia and Kenya. Int J Pest Manage 65: 1\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKutka F (2011) Open-Pollinated vs. Hybrid Maize Cultivars. Sustainability 3: 1531\u0026ndash;1554.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLee J, Chin JH, Ahn SN, Koh HJ (2015) Brief history and perspectives on plant breeding. In: Koh KH, Kwon SY, Thomson M (ed) Current Technologies in Plant Molecular Breeding: A Guide Book of Plant Molecular Breeding for Researchers, Springer, Dordrecht, pp. 1\u0026ndash;14.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLenth R (2023) emmeans: Estimated Marginal Means, aka Least-Squares Means_. R package version 1.8.5, \u0026lt;\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://CRAN.R-project.org/package=emmeans\u0026gt;\u003c/span\u003e\u003cspan address=\"https://CRAN.R-project.org/package=emmeans%3E\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLiebman M, Davis AS (2009) Managing weeds in organic farming systems: An ecological approach. In: Francis C (ed) Organic Farming: The Ecological System. Agronomy Monograph, Madison, WI, USA, pp 173\u0026ndash;195.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMacRobert JF, Setimela PS, Gethi J, Worku M (2014) Maize Hybrid Seed Production Manual. Mexico, D.F. CIMMYT.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMcCullagh P, Nelder JA (1989) Generalized Linear Models. London: Chapman and Hall. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1007/978-1-4899-3242-6\u003c/span\u003e\u003cspan address=\"10.1007/978-1-4899-3242-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeagher RL, Nagoshi RN, Stuhl CJ (2011) Oviposition Choice of Two Fall Armyworm (Lepidoptera: Noctuidae) Host Strains. J Insect Behav 24: 337\u0026ndash;347.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMontezano DG, Specht A, Sosa-G\u0026oacute;mez DR, Roque-Specht VF, Sousa-Silva JC, Paula-Moraes SD, Hunt TE (2018) Host plants of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) in the Americas. Afr Entomol 26: 286\u0026ndash;300.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMutyambai DM, Niassy S, Calatayud PA, Subramanian S (2022) Agronomic Factors Influencing Fall Armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e) Infestation and Damage and Its Co-occurrence with Stemborers in Maize Cropping Systems in Kenya. Insects 13: 266. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/insects13030266\u003c/span\u003e\u003cspan address=\"10.3390/insects13030266\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNagoshi RN, Meagher RL, Hay-Roe M (2012) Inferring the annual migration patterns of fall armyworm (Lepidoptera:Noctuidae) in the United States from mitochondrial haplotypes. Ecol Evol 2: 1458\u0026ndash;1467.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOdeyemi OO, Lawal BO, Owolade OF, Olasoji JO, Egbetokun OA, Oloyede- Kamiyo QO, Omodele T, Anjorin FB (2020) Fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J.E. Smith) (Lepidoptera: Noctuidae): threat to maize production in Nigeria. Nig Agric J 51: 101\u0026ndash;108.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOgunfunmilayo AO, Kazeem SA, Idoko JE, Adebayo RA, Fayemi EY, Adedibu OB, Ofuya TI (2021) Occurrence of natural enemies of fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) in Nigeria. Plos one 16: 254\u0026ndash;328.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOjumoola AO, Omoloye AA (2023) Agroecological zones influence maize infestation and damage severity by the fall armyworm \u003cem\u003e(Spodoptera frugiperd\u003c/em\u003ea (J. E. Smith)) in southwestern Nigeria Acta Agric 119: 1\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOlaniyan AB (2015) Maize: Panacea for hunger in Nigeria. Afr J Plant Sci 3: 155\u0026ndash;174.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePiepho HP (2004) An algorithm for a letter-based representation of all pairwise comparisons, J Comput Graph Stat 13: 456\u0026ndash;466.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePiesik D, Rochat D, Delaney KJ, Marion-Poll F (2012). Orientation of European corn borer first instar larvae to synthetic green leaf volatiles. J Applied \u0026Egrave;ntomol 137: 234\u0026ndash;240.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePogue MG (2002) A world revision of the genus \u003cem\u003eSpodoptera guen\u0026eacute;e\u003c/em\u003e (Lepidoptera: Noctuidae). Mem Am Entomol Soc 43:1\u0026ndash;202.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eProwell DP, McMichael M, Silvain JF (2004) Multilocus genetic analysis of host use, introgression, and speciation in host strains of fall armyworm (Lepidoptera: Noctuidae). Ann Entomol Soc Am 97:1034\u0026ndash;1044.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePӧykkӧ H (2006) Females and larvae of a geometrid moth, \u003cem\u003eCleorodes lichenaria\u003c/em\u003e, prefer a lichen host that assures shortest larval period. Environ Entomol 35:1669\u0026ndash;1675.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eR Core Team (2022) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.R-project.org/\u003c/span\u003e\u003cspan address=\"https://www.R-project.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRefsnider JM, Janzen FJ (2010). Putting eggs in one basket: Ecological and evolutionary hypotheses for variation in oviposition-site choice. Annu Rev Ecol Evol Syst 41: 39\u0026ndash;57.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRojas JC, Kolomiets MV, Bernal JS (2018) Nonsensical choices? Fall armyworm moths choose seemingly best or worst hosts for their larvae, but neonate larvae make their own choices. PloS one 13(5): e0197628.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRojas JC, Virgen A and Cruz-Lopez L (2003) Chemical and tactile cues influencing oviposition of a generalist moth, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae). Environ Entomol 32: 1386\u0026ndash;1392.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRussianzi W, Anwar R, Triwidodo H (2021) Biostatistics of fall armyworm \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e in maize plants in Bogor, West Java, Indonesia. Biodiversitas 22: 3463\u0026ndash;3469.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRwomushana I, Bateman M, Beale T, Beseh P, Cameron K., Chiluba M, Tambo J (2018) Fall armyworm: impacts and implications for Africa. Wallingford, UK\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSaeed S, Sayyed AH, Ahmad I (2010) Effect of host plants on life-history traits of \u003cem\u003eSpodoptera exigua\u003c/em\u003e (Lepidoptera: Noctuidae). J Pest Sci 83: 165\u0026ndash;172.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSakamoto Y, Ishiguro M, Kitagawa G (1986) \u003cem\u003eAkaike Information Criterion Statistics\u003c/em\u003e. D. Reidel Publishing Company.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSingh GM, Xu J, Schaefer D, Day R, Wang Z, Zhang F (2021) Maize Diversity for Fall Armyworm Resistance in a Warming World. Crop Sci 62: 1\u0026ndash;19.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSisay B, Sevgan S, Weldon CW, Kr\u0026uuml;ger K, Torto B, Tamiru A. (2023) Responses of the fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e) to different host plants: Implications for its management strategy. Pest Manage Sci 79: 845\u0026ndash;856.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSobhy IS, Tamiru A, Chiriboga MX, Nyagol D, Cheruiyot D, Chidawanyika F, Subramanian S, Midega CAO, Bruce TJA, Khan ZR (2022) Bioactive Volatiles from Push-Pull Companion Crops Repel Fall Armyworm and Attract Its Parasitoids. Front Ecol Evol 10:883020. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3389/fevo.2022.883020\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2022.883020\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSokame BM, Subramanian S, Kilalo DC, Juma G, Calatayud PA (2020) Larval dispersal of the invasive fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e, the exotic stemborer \u003cem\u003eChilo partellus\u003c/em\u003e, and indigenous maize stemborers in Africa. Entomol Exp Appl, 168: 322\u0026ndash;331.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSoteloCardona P, Chuang WP, Lin MY, Chiang MY, Ramasamy S (2021) Oviposition preference not necessarily predicts offspring performance in the fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) on vegetable crops. Sci Rep 11:15885; \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/s41598-021-95399-4\u003c/span\u003e\u003cspan address=\"10.1038/s41598-021-95399-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSparks AN (1979) A review of the biology of the fall armyworm. Fla Entomol 62: 82e87. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.2307/3494083\u003c/span\u003e\u003cspan address=\"10.2307/3494083\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSzendrei Z, Rodriguez-Saona C (2010) A meta-analysis of insect pest behavioral manipulation with plant volatiles. Entomol Exp Appl 134: 201\u0026ndash;210.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTambo JA, Kansiime MK, Mugambi I, Agboyi LK, Beseh PK, Day R (2023) Economic impacts and management of fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e) in smallholder agriculture: a panel data analysis for Ghana. CABI Agric Biosci 4:38 \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s43170-023-00181-3\u003c/span\u003e\u003cspan address=\"10.1186/s43170-023-00181-3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTiwari S (2022) Host plant preferences by fall armyworm \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (JE Smith) (Lepidoptera: Noctuidae) on the range of potential host plant species. J Agric For 5: 25\u0026ndash;33.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTogola A, Meseka S, Menkir A, Badu-Apraku B, Boukar O, Tam\u0026ograve; M, Djouaka R (2018) Measurement of Pesticide Residues from Chemical Control of the invasive \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) in a Maize Experimental Field in Mokwa, Nigeria. Int J Environ Res Public Health 15: 849\u0026ndash;860.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eUnited States Department of Agriculture (USDA) (2020). World Agricultural Production: Global Market Analysis, International Production Assessment Division, Washington, DC \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://apps.fas.usda.gov/psdonline/circulars/production.pdf\u003c/span\u003e\u003cspan address=\"https://apps.fas.usda.gov/psdonline/circulars/production.pdf\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVolp TV, Zalucki MP, Furlong MJ (2022). What Defines a Host? Oviposition Behavior and Larval Performance of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) on Five Putative Host Plants, J Econ Entomol 115: 1744\u0026ndash;1751.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang L, Yang A, He C, Qu M, Zhang J (2008). Creation of new maizegermplasm using alien introgression from \u003cem\u003eZea mays\u003c/em\u003e ssp. mexicana. Euphytica 164: 789\u0026ndash;801.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYactayo-Chang JP, Mendoza J, Willams SD, Rering CC, Beck JJ, Block AK (2021). \u003cem\u003eZea mays\u003c/em\u003e Volatiles that Influence Oviposition and Feeding Behaviors of \u003cem\u003eSpodoptera frugiperda.\u003c/em\u003e J Chem Ecol 47: 799\u0026ndash;809.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang X, Wyckhuys KA, Jia X, Nie F, Wu K. (2021). Fall armyworm invasion heightens pesticide expenditure among chinese smallholder farmers. J Environ Manage, 282, 111949.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang G, Espelie KE, Wiseman BR, Ishenhour DJ (1993a) Effects of corn foliar cuticular lipids on the movement of fall armyworm (Lepidoptera: Noctuidae) neonate larvae. Fla Entomol 76: 302\u0026ndash;316\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang G, Wiseman BR, Ishenhour DJ, Espelie KE (1993b) Chemical and ultrastructural analysis of corn cuticular lipidsand their effects on fall armyworm larva. J Chem Ecol 9: 2055\u0026ndash;2074.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang J, Ma C, Jia R, Zhang H, Zhao Y, Yue H, Li H, Jiang X (2023) Different responses of two maize cultivars to \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) larvae infestation provide insights into their differences in resistance. Front Plant Sci 14:1065891. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3389/fpls.2023.1065891\u003c/span\u003e\u003cspan address=\"10.3389/fpls.2023.1065891\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang QY, Zhang YL, Quandahor, P., Gou, Y.P., Li, C.C., Zhang, K.X., Liu, C.Z. (2023). Oviposition Preference and Age-Stage, Two-Sex Life Table Analysis of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (Lepidoptera: Noctuidae) on different Maize Varieties. Insects 14: 413. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/insects14050413\u003c/span\u003e\u003cspan address=\"10.3390/insects14050413\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Fall armyworm, maize plant volatiles, multiple choice experiment, olfactometer bioassay, resistant varieties","lastPublishedDoi":"10.21203/rs.3.rs-4601270/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4601270/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003ePhytophagous insects likely select suitable host plants for oviposition based on olfactory and tactile cues. However, details of how insects differentiate among different plant varieties are often unclear. The fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e J. E. Smith) is a highly destructive pest on maize, but little is known about the attraction and oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e to different maize varieties, particularly in the context of sub-Saharan Africa, where the insect is a major threat to maize production. We determined the oviposition preference of \u003cem\u003eS. frugiperda\u003c/em\u003e females on six different maize plant varieties three of which were hybrid varieties and three were open pollinated varieties, in multiple-choice and no-choice assays. We also evaluated the attraction preference of \u003cem\u003eS. frugiperda\u003c/em\u003e larvae on these maize varieties, using an olfactometer bioassay. We found that \u003cem\u003eS. frugiperda\u003c/em\u003e females oviposited significantly less egg masses on the hybrid varieties \u003cem\u003eDEKAIB\u003c/em\u003e and \u003cem\u003e30Y87\u003c/em\u003e than on the other varieties tested, and that females oviposited less on the hybrid maize varieties compared to the open pollinated maize varieties overall. Additionally, we found that \u003cem\u003eS. frugiperda\u003c/em\u003e larvae were more attracted to the open pollinated variety LMFP than to clean air, which was not the case for any of the other maize varieties tested. Taken together, our results show that \u003cem\u003eS. frugiperda\u003c/em\u003e responds differentially to the different maize varieties and that hybrid maize varieties seem less attractive. Further investigating the chemistry of hybrid maize varieties like \u003cem\u003eDEKAIB\u003c/em\u003e might yield clues on how to breed maize varieties with increased resistance against \u003cem\u003eS. frugiperda\u003c/em\u003e infestation.\u003c/p\u003e","manuscriptTitle":"Oviposition behavior and larval attraction of the fall armyworm Spodoptera frugiperda to different maize plant varieties","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-12 15:14:03","doi":"10.21203/rs.3.rs-4601270/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"59b7d9cd-288d-4606-ab43-28b88f0d18b5","owner":[],"postedDate":"July 12th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-07-15T12:01:41+00:00","versionOfRecord":[],"versionCreatedAt":"2024-07-12 15:14:03","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4601270","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4601270","identity":"rs-4601270","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}