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Article

Love in the Time of Pyrethroids: Mating Behavior of Sitophilus zeamais Is Influenced by Sublethal Concentrations of λ-Cyhalothrin and Lateralization

by
Maria C. Boukouvala
,
Nickolas G. Kavallieratos
*,
Demeter Lorentha S. Gidari
,
Constantin S. Filintas
,
Anna Skourti
,
Vasiliki Panagiota C. Kyrpislidi
and
Dionysios P. Skordos
Laboratory of Agricultural Zoology and Entomology, Faculty of Crop Science, Agricultural University of Athens, 75 Iera Odos Str., 11855 Athens, Greece
*
Author to whom correspondence should be addressed.
Insects 2025, 16(8), 865; https://doi.org/10.3390/insects16080865
Submission received: 23 July 2025 / Revised: 8 August 2025 / Accepted: 19 August 2025 / Published: 20 August 2025
(This article belongs to the Special Issue Integrated Pest Management in Agricultural Crops and Forest Ecosystems—2nd Edition)
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Simple Summary

The present study investigates the alterations of the mating behavioral traits of Sitophilus zeamais under sublethal concentrations of the pyrethroid λ-cyhalothrin and the impact of lateralization. The treated individuals demonstrated altered behavior compared to the untreated individuals. Significant changes were observed in the mating success rates, the mate recognition time, and the duration of the copulation, among the tested groups (control, LC10, and LC30). The lateralization significantly impacted mating behavioral traits, such as the interaction between the males and females and the duration of the copulation. These results suggest that low concentrations of λ-cyhalothrin might disrupt the reproduction of S. zeamais, offering a new angle for pest control strategies that focus on behavioral alterations.

Abstract

Sitophilus zeamais Motschulsky (Coleoptera: Curculionidae) is one of the most destructive pests of stored grains worldwide. Sublethal concentrations of insecticides are known to influence insect behavior, potentially disrupting critical processes such as mating. This study investigated the effects of λ-cyhalothrin at the lethal concentration (LC) values LC10 and LC30 and lateralization on the mating behavior patterns of S. zeamais males. Results showed that the exposure to sublethal concentrations of λ-cyhalothrin significantly altered the copulation success rate and key time-related parameters, including mate recognition and copulation duration, while the lateralization caused significant differences in mating time-related parameters within each tested group (control, LC10, and LC30). Additionally, the λ-cyhalothrin-treated groups showed prolonged mate recognition times and required more mounting attempts to achieve mating. These findings highlight the potential of sublethal insecticide applications to control S. zeamais populations by impairing reproduction.

1. Introduction

Sitophilus zeamais Motschulsky (Coleoptera: Curculionidae) is a noxious insect of stored grains worldwide, particularly in tropical and subtropical regions [1,2,3,4]. It is a primary pest of maize [5,6] but is also known to infest other grains such as wheat, rice, and sorghum [7,8,9]. In addition to direct consumption of the grains, S. zeamais infestation leads to increased moisture content and fungal contamination, further compounding post-harvest losses [10,11].
The type II pyrethroid λ-cyhalothrin is an efficient insecticidal agent against various insect species, including stored-product pests [12,13,14,15,16]. It interacts with neuronal membrane sodium (Na+) channels, which are essential for the response of the insect to nerve inputs. When λ-cyhalothrin reaches the active site, it causes uncontrollable nerve discharges, potentially leading to tremors, twitches, and paralysis [17]. It can also interact with the calcium and chloride channels, quickly impairing muscular control and feeding disruptions, as it is easily absorbed by biological tissues and penetrates the cuticle of the insect [18]. λ-Cyhalothrin has also been tested for the effect of the sublethal concentrations on numerous insect species [19,20,21,22]. Under the influence of sublethal insecticidal concentrations, insects often exhibit various behavioral alterations, including locomotion, selection of oviposition sites, mating behavior and copulation success, and disruptions in the chemical communication systems [21,23,24,25].
Lateralization (i.e., asymmetries in the two hemispheres of the brain and behavior) has been thoroughly investigated in vertebrates, ants, and honeybees [26,27,28,29,30,31]. In recent years, various studies have explored the lateralization of the mating behavior of stored product pests [24,32,33,34].
The rapid population growth of S. zeamais favors the extensive damage of the infested commodities [35,36,37,38]. It has been previously documented that insecticides at sublethal concentrations compromise the development of stored-product insects [39,40]. However, there is no data about the effect of λ-cyhalothrin at low concentrations on the mating behavior of S. zeamais. Thus, the current study aims to investigate the impact of λ-cyhalothrin LC10 and LC30 on the courtship and mating of S. zeamais, taking into account the influence of males’ lateralization.

2. Materials and Methods

2.1. Sitophilus zeamais Colonies and Sex Recognition

The male and female S. zeamais adults that were used in the bioassays were obtained from the Laboratory of Agricultural Zoology and Entomology (Athens, Greece). Sitophilus zeamais individuals were reared on insecticide-free maize in devoid of infestation or impurities [41]. The colonies were maintained in controlled conditions (65% relative humidity (RH) and 30 °C) and continuous darkness [21,42]. One day, newly emerged adults, obtained from separately kept maize kernels, were sexed according to Halstead [43] and Tolpo and Morrison [44], based on the pattern of their rostrum punctuation, length, and texture. Then they were kept separately, by sex, in jars with rearing medium until sexual maturation (4 days old) [36]. The female individuals were marked with white color on their thorax to allow sex recognition during the bioassays.

2.2. Insecticide

The commercial formulation ICON 10 CS (Syngenta, Anthousa, Greece), with λ-cyhalothrin as the active ingredient (a.i.) at 10.7 w/w, was used for the bioassays.

2.3. Contact Toxicity Bioassays

To determine the LC10, LC30, and LC50 of λ-cyhalothrin against male and female adults of S. zeamais, six dilutions (i.e., 0.0025, 0.0013, 0.0008, 0.000625, 0.0005, and 0.00042 mg a.i./cm2) were prepared in distilled water. Then, separate filter papers (Whatman No. 1) cut into a circular shape and placed at the bottom of a Petri dish (50.27 cm2) were impregnated, using a micropipette, with 1 mL of the various dilutions and left to dry at 30 °C for 2 h. Control filter papers were impregnated with 1 mL of distilled water. After the filter papers had dried, 20 S. zeamais individuals were placed in each dish (6 concentrations × 5 replications = 30 dishes). Then the dishes were maintained in 65% RH, 30 °C, and continuous darkness for 72 h, and the total number of dead adults was counted. Prior to the behavioral bioassays, new male and female S. zeamais individuals were exposed to the LC10, LC30 of λ-cyhalothrin or the control, as described above.

2.4. Mating Behavior Bioassays

The experiment was conducted in a Petri dish arena surrounded by a filter paper box to prevent any disruption of the individuals from the observer or the surrounding area [32]. The walls of the dishes were covered with liquid polytetrafluoroethylene (Sigma-Aldrich Chemie GmbH, Taufkirchen, Germany) to prevent insects’ escapes. Prior to each observation, S. zeamais individuals were adjusted to the natural conditions of light for a 3 h interval [24,33,34,45]. A male and a female S. zeamais (virgins), treated with LC10 or LC30 of λ-cyhalothrin or the control, were placed in the arena and were visually recorded for 60 min (min) or until the end of the mating process [46]. For each mating pair, the following parameters were recorded: (1) the side (back, left, right or front) of the body of the female that the male approached, (2) the mounted side (back, left, right or front) of the body of the female, (3) the mounting attempts of males (number), (4) the stroking behavior, if any, (5) the copulation success, if any, and (6) the side (back, left, right or front) of leaving (which side the male leaves the body of the female). The courtship time parameters were observed for (1) the duration of mate recognition, and (2) the duration of the interaction between the two individuals, mounting attempts of males, and the walking that the individuals performed attached, consisting of precopula. The duration of the copulation was also recorded. When 50 successful mating pairs were observed per treatment or control, counting was completed. Thus, 90, 131, and 166 pairs were recorded in total for control, LC10, and LC30, respectively. However, pairs where females stayed next to the arena walls, affecting the males’ approach choice, and pairs that were motionless or did not engage in any sexual contact were rejected [33]. Therefore, 13, 20, and 27 pairs were removed from the control, LC10, and LC30 groups, respectively. As a result, there were 77 (control), 111 (LC10), and 139 (LC30) mating pairs included in the statistical analysis.

2.5. Statistical Analysis

LC10, LC30, and LC50 of λ-cyhalothrin were estimated using probit analysis by the R software (version 2.15.1), with a 95% confidence interval (CI) [47,48]. Data on the laterality behavior was analyzed using the software JMP 16.2 [49]. The courtship time parameters, which did not follow a normal distribution, were analyzed using the Steel–Dwass test with a significance level of 0.05 [45]. The mating success was calculated by a weighted generalized linear model with binomial distribution: y = Xβ + ε (y: the vector of observations, i.e., successful or not successful copulation; X: the incidence matrix; β: the vector of fixed effect, i.e., λ-cyhalothrin LC10-, LC30-exposed, and control; ε: the vector of the random residual effect). To detect the significant differences between values, the 0.05 level of probability was used.

3. Results

3.1. Contact Toxicity of λ-Cyhalothrin on S. zeamais Adults

The bioassays on the contact toxicity of λ-cyhalothrin on S. zeamais adults are presented in Table 1.

3.2. Impact of λ-Cyhalothrin and Laterality on the Mating Success of S. zeamais

Most of the control males showed a left-biased approach tendency (48.1 out of 100%), compared to right and back (37.6 and 14.3 out of 100%, respectively). The majority of the left-biased control males mounted their mate from their left side (37.7% out of 48.1%). The 12.8 out of 37.7% left-biased males performed stroking behavior, of which 6.6% (out of 12.8%) mated successfully. The 24.9% out of 37.7% did not perform stroking behavior, where 18.3% out of 24.9% achieved successful copulations and left the body of the females from their back, left, and right sides at rates of 7.5, 9.2, and 1.6%, respectively (Figure 1).
Concerning the λ-cyhalothrin LC10-treated S. zeamais individuals, most of the males had a left-biased tendency (42.4 out of 100%), compared to the other directions (36.3, 16.2, and 5.1 out of 100% for right-biased, back-side, and front-side, respectively). The 32.5 out of 42.4% of the left-biased males tend to mount the females from their left side. Most of them, 27.1 out of 32.5%, did not perform stroking behavior, where 15.4 out of 27.1% of these males achieved successful copulations, leaving the body of the female from their back, left, and right sides at rates of 8.2, 5.4, and 1.8%, respectively. In the case of right-biased approaches (36.3%), most of the males performed mountings from the right side of the female’s body (29.8 out of 36.3%), with 26.1 out of 29.8% of them not showing stroking behavior, and finally, 7.5% (out of 26.1%) mated with success (Figure 2).
A similar pattern, as in the control and the λ-cyhalothrin LC10 groups, was recorded for the λ-cyhalothrin LC30-treated S. zeamais male individuals, showing a left-biased tendency for the approach (41.7 out of 100%), compared to right-biased, back-side, and front-side (33.1, 15.1, and 10.1 out of 100%, respectively). Most of the left-biased λ-cyhalothrin LC30-treated males mounted the females from their left side (35.3 out of 41.7%). The 29.5 out of 35.3% did not perform stroking behavior, where 12.2 out of 29.5% of males had successful copulation and 17.3 out of 29.5% failed to mate. The majority of males that demonstrated right-biased approaches mounted females from their right side (28.8 out of 33.1%), did not perform the stroking behavior (25.9 out of 28.8%), while only 7.2 out of 25.9% succeeded in copulation (Figure 3).
In Figure 4, the mating success of the treated (LC10, LC30), and control S. zeamais is presented. Fifty mating couples of each tested group achieved successful copulations. The successful copulation of males was affected by their exposure to LC10 and LC30 of λ-cyhalothrin (χ2 = 29.0, df = 2, p < 0.01). Mating success of S. zeamais males exposed to control and LC30 of λ-cyhalothrin was significantly affected (χ2 = 6.9, df = 1, p > 0.01 and χ2 = 11.0, df = 1, p > 0.01, respectively). Mating success of λ-cyhalothrin LC10-treated males was not affected significantly (χ2 = 1.1, df = 1, p > 0.05) (Figure 4).

3.3. Impact of λ-Cyhalothrin and Laterality on S. zeamais Mating Time Parameters

3.3.1. Impact of the Direction of the Approach

The side that males approached the females affected the mate recognition time. Within each tested group (control, λ-cyhalothrin LC10, and λ-cyhalothrin LC30), there were significant differences among the various directions. Significant differences were also observed among the tested groups. The λ-cyhalothrin LC30-treated males, which approached their mates from their back side, performed a significantly longer time interval to detect their mates (3.9 min), compared to all the other males of the three tested groups (control, λ-cyhalothrin LC10, and λ-cyhalothrin LC30). The significantly shortest mate recognition time was recorded for the left-biased control males (1.8 min), compared to all the other males of the different directions and tested groups (Table 2).
The intervals of the interaction between the two individuals (female and male) were significantly influenced by the direction of the approach and the insecticidal treatment. The left-biased control males spent significantly the shortest time (3.8 min) interacting with the females, compared to those of the control group that approached from the back and right sides, as well as compared to the males of all directions of the two λ-cyhalothrin-treated groups (Table 2).
The control males that first approached their mates from their back side performed significantly longer walking time (7.2 min), compared to all the other males (left- and right-biased control males, males of the λ-cyhalothrin LC10 and LC30 groups that first approached their mates from their back, left, right, or front side) (Table 2).
Concerning the mounting attempts, control males performed significantly fewer attempts to mount their mates (1.6 and 1.7 times for the left- or right- and for the back- side approaching males, respectively), compared to the λ-cyhalothrin LC10 or the λ-cyhalothrin LC30 males. λ-Cyhalothrin LC30 males that first approached their mates from their front side performed significantly more mounting attempts, vs. all the other males, except for the front-approaching males of the λ-cyhalothrin LC10 group (Table 2).
Regarding the duration of the copulation, males of all directions of the control group mated for significantly longer times (276.9, 274.1, and 268.1 min, for the left-, right-, and back-side approaching males), compared to the males of all directions of the two λ-cyhalothrin-treated groups. The front-side approaching males of the λ-cyhalothrin LC30 group copulated for significantly shorter time (18.6 min), compared to all the other males of all directions and the tested group (Table 2).

3.3.2. Impact of the Mounting Direction

The mate recognition time was not significantly affected by the laterality within the control group; however, it was significantly increased in the λ-cyhalothrin-treated groups, compared to the control. The significantly shortest mate recognition times were recorded for the control males (2.1, 2.0, and 1.9 min for the back-, right-, and left-mounting males). The longest time (3.9 min) was detected for the λ-cyhalothrin LC30-treated males that mounted their mates from their back side, compared to all the other males, except for the front-mounting males of the two λ-cyhalothrin groups (Table 3).
The time of the interaction was significantly longer for the λ-cyhalothrin LC30 males that mounted their mates from their front side (9.9 min), compared to all the other males of all directions of the three tested groups (control, λ-cyhalothrin LC10, and λ-cyhalothrin LC30), except for the front-mounting males of the λ-cyhalothrin LC10 group (8.9 min). In contrast, the shortest interaction time (4.5 min) was recorded at the left-mounting males of the control group, significantly shorter compared to all the other males (Table 3).
The walking time was not significantly influenced by the lateralization within each tested group (control, λ-cyhalothrin LC10, and λ-cyhalothrin LC30) but was significantly influenced by the insecticidal application. The longest walking periods were detected for the control males (5.3 and 5.2 min for the back- or left- and right-mounting males, respectively), which were significantly different from the λ-cyhalothrin LC10- and λ-cyhalothrin LC30-treated males (Table 3).
The number of mounting attempts for the control males of all directions was significantly shorter (1.7 and 1.6 times for the back- or left- and right-mounting males, respectively), compared to all the males of the λ-cyhalothrin-treated groups (Table 3).
The application of the insecticide and the laterality significantly affected the duration of the copulation of each tested group (control, λ-cyhalothrin LC10, and λ-cyhalothrin LC30). The use of λ-cyhalothrin LC10 and LC30 caused a significant decrease in the copulation time, compared to the control group. The right-mounting males of the control group copulated for significantly longer time (309.7 min), compared to all the other males of all directions and tested groups. The front-mounting males of the λ-cyhalothrin LC10 group and the right- and front-mounting males of the λ-cyhalothrin LC30 group performed the shortest durations of copulation (61.8, 51.8, and 44.6 min, respectively) (Table 3).

4. Discussion

The current study demonstrates that λ-cyhalothrin, even at low concentrations, radically alters key aspects of mating interactions, including copulation success, and time-related parameters such as mate recognition time and copulation duration. The observed alterations in the mating traits at both λ-cyhalothrin-treated groups (LC10 and LC30) suggest neurophysiological interference driven by the insecticidal mode of action on the neural ion channels of the insects [17,18], given that the behavioral traits refer to the function of the brain [50]. The influence of the insecticidal action proved decisive for the success of copulation. The present study documents that the overall successful copulation rate followed a decreasing order among treatments, i.e., LC30 (41.2%) < LC10 (51.6%) < control (65.3%). Similarly, when Alphitobius diaperinus (Panzer) (Coleoptera: Tenebrionidae) adults were exposed to sublethal concentrations of another pyrethroid insecticide, α-cypermethrin, the successful copulation rates were 87.2, 63.4, and 64.4% for the control, LC10, and LC30 groups, respectively [24].
Here, the mate detection time was significantly increased in the insecticide-treated groups. This pattern has also been observed in other studies. Prostephanus truncatus (Horn) (Coleoptera: Bostrychidae) adults exposed to sublethal concentrations of Acmella oleracea (L.) R.K. Jansen (Asteraceae) hexane extract resulted in prolonged mate recognition duration [51]. Additionally, A. diaperinus α-cypermethrin LC10- and LC30-treated adults resulted in increased mate detection duration [24]. The number of mounting attempts was also notably higher in all the side tendences of the λ-cyhalothrin LC10 and LC30 groups, vs. the control. Given that the mounting is crucial for the mating of S. zeamais [46,52], the predicament of this behavioral trait could prevent the achievement of successful copulation. Insects depend on mating as a means of population growth, establishment in new habitats, and coexistence with other species [53]. Since the difficulty or delay of mate recognition, decreased copulation time, and mating success rates have been proven to lead to the inability of the population growth [54,55], the sublethal concentrations of λ-cyhalothrin could decrease the population of S. zeamais.
Sitophilus zeamais females create holes in the kernel, where they lay eggs and then seal the holes using a secretion [56,57]. Larvae borrow the internal part of the kernels, which contain carbohydrates and other nutrients, causing severe weight loss [57,58]. Given that the development until the adult stage takes place inside the kernel, the detection of the infestation and the control of this species is challenging [59]. Thus, it is important to target the management of S. zeamais adults. Assuming that the negative effect of the sublethal concentrations of λ-cyhalothrin on the mating behavior and success of S. zeamais could prevent the fecundity and the oviposition process, the current treatment could eliminate the qualitative and quantitative damage of the maize kernels. Interestingly, a recent study on another coleopteran species demonstrated that sublethal concentrations of λ-cyhalothrin significantly reduced the average fecundity and prolonged the durations of all the developmental stages of Henosepilachna vigintioctomaculata (Coccinellidae) [16]. However, this issue needs to be further investigated.
Left-biased males of all three tested groups achieved higher copulation rates compared to those of the other sides (back, right, and front). This pattern is common among various coleopteran families. For instance, it has been evident that different tenebrionids tend to achieve higher rates of successful copulations when they demonstrate a left-biased tendency [33,34]. The exposure of S. zeamais adults to λ-cyhalothrin did not affect the tendency of the side of the approach. Similarly, A. diaperinus males and females, that had been treated with α-cypermethrin LC10 and LC30, retained their right-biased tendency approaching their mates, as in the control group [24]. Furthermore, when Romano et al. [32] tested the impact of the geographical origin and the rearing medium of Sitophilus oryzae (L.) (Coleoptera: Curculionidae), the left-biased copulation tendency, which led to higher mating success, was not affected for the adults of all three strains (two Greek strains, one reared on wheat and one on maize, and a Peruvian strain reared on maize). The maintenance of specific lateral tendencies under the influence of insecticidal agents or environmental parameters indicates that they are strongly stable. Whether they can be altered under stronger influences, such as higher insecticidal concentrations, needs further research.

5. Conclusions

The results of the current study highlight the importance of incorporating behavioral traits into the assessment of sublethal insecticidal impacts. Sublethal concentrations of λ-cyhalothrin significantly disrupted the mating behavior of S. zeamais, introducing one other component in the concept of management strategies of this species. Nevertheless, future studies employing metabolomics, i.e., studying the changes in the metabolic paths, could shed further light on how λ-cyhalothrin alters the mating traits in stored-product insects.

Author Contributions

Conceptualization, M.C.B., N.G.K., D.L.S.G., C.S.F. and A.S.; methodology, M.C.B., N.G.K., D.L.S.G., C.S.F. and A.S.; software, M.C.B., N.G.K. and D.L.S.G.; validation, M.C.B., N.G.K., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; formal analysis, M.C.B., N.G.K. and D.L.S.G.; investigation, M.C.B., N.G.K., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; resources, N.G.K.; data curation, M.C.B., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; writing—original draft preparation, M.C.B., N.G.K., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; writing—review and editing, M.C.B., N.G.K., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; visualization, M.C.B., N.G.K., D.L.S.G., C.S.F., A.S., V.P.C.K. and D.P.S.; supervision, N.G.K.; project administration, N.G.K.; funding acquisition, N.G.K. All authors have read and agreed to the published version of the manuscript.

Funding

The project 80375 (Agricultural University of Athens, Account for Research Funds) supported this study partially.

Data Availability Statement

Data is available within the article.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Banga, K.S.; Kumar, S.; Kotwaliwale, N.; Mohapatra, D. Major insects of stored food grains. Int. J. Chem. Stud. 2020, 8, 2380–2384. [Google Scholar] [CrossRef]
  2. Stathers, T.; Holcroft, D.K.L.; Mvumi, B.; English, A.; Omotilewa, O.; Kocher, M.; Ault, J.; Torero, M. A scoping review of interventions for crop postharvest loss reduction in sub-Saharan Africa and South Asia. Nat. Sustain. 2020, 3, 821–835. [Google Scholar] [CrossRef]
  3. Srivastava, S.; Mishra, H.N. Development of microencapsulated vegetable oil powder based cookies and study of its physicochemical properties and storage stability. LWT 2021, 152, 112364. [Google Scholar] [CrossRef]
  4. Achimón, F.; Peschiutta, M.L.; Brito, V.D.; Beato, M.; Pizzolitto, R.P.; Zygadlo, J.A.; Zunino, M.P. Exploring contact toxicity of essential oils against Sitophilus zeamais through a meta-analysis approach. Plants 2022, 11, 3070. [Google Scholar] [CrossRef]
  5. Erenso, T.F.; Berhe, D.H. Effect of neem leaf and seed powders against adult maize weevil (Sitophilus zeamais Motschulsky) mortality. Agric. Res. 2016, 2, 90–94. [Google Scholar] [CrossRef]
  6. Brito, V.D.; Achimón, F.; Pizzolitto, R.P.; Ramírez Sánchez, A.; Gómez Torres, E.A.; Zygadlo, J.A.; Zunino, M.P. An alternative to reduce the use of the synthetic insecticide against the maize weevil Sitophilus zeamais through the synergistic action of Pimenta racemosa and Citrus sinensis essential oils with chlorpyrifos. J. Pest Sci. 2021, 94, 409–421. [Google Scholar] [CrossRef]
  7. Carvalho, M.O.; Fradinho, P.; Martins, M.J.; Magro, A.; Raymundo, A.; Sousa, I. Paddy rice stored under hermetic conditions: The effect of relative humidity, temperature and storage time in suppressing Sitophilus zeamais and impact on rice quality. J. Stored Prod. Res. 2019, 80, 21–27. [Google Scholar] [CrossRef]
  8. Zhou, Y.; Hui, Y.B.; Feng, L.F.; Zhou, T.; Wang, Q. A method for reconstructing the internal morphological structure of wheat kernels upon Sitophilus zeamais infestation. J. Stored Prod. Res. 2020, 88, 101676. [Google Scholar] [CrossRef]
  9. Suleiman, M.; Ibrahim, N.D.; Rugumamu, C.P. Application of some botanicals in the protection of sorghum grains against the maize weevil, Sitophilus zeamais Motsch. (Coleoptera: Curculionidae). Proc. Zool. Soc. 2021, 74, 219–226. [Google Scholar] [CrossRef]
  10. Bhusal, K.; Khanal, D. Role of maize weevil, Sitophilus zeamais Motsch. on spread of Aspergillus section flavi in different Nepalese maize varieties. Adv. Agric. 2019, 2019, 7584056. [Google Scholar] [CrossRef]
  11. Sebayang, A.; Rubiana, R.; Sipi, S.; Manwan, S.W.; Fattah, A.; Arrahman, A.; Saenong, M.S. Sitophilus zeamais (Motschulsky): The primary obstacles in the maize quality and quantity. IOP Conf. Ser. Earth Environ. Sci. 2023, 1230, 12089. [Google Scholar] [CrossRef]
  12. Shakoori, F.R.; Feroz, A.; Gondal, A.; Akram, S.; Riaz, T. Impact of λ-cyhalothrin on carbohydrate metabolizing enzymes and macromolecules of a stored grain pest, Trogoderma granarium. Pak. J. Zool. 2018, 50, 1467–1474. [Google Scholar] [CrossRef]
  13. Li, X.R.; Li, Y.; Wang, W.; He, N.; Tan, X.L.; Yang, X.Q. LC50 of lambda-cyhalothrin stimulates reproduction on the moth Mythimna separata (Walker). Pestic. Biochem. Phys. 2019, 153, 47–54. [Google Scholar] [CrossRef]
  14. Keyhanian, A.A.; Barari, H.; Mobasheri, M.T. Comparison of the efficacy of insecticides, alphacypermethrin and lambda-cyhalothrin, against canola flea beetles. Appl. Entomol. Phytopathol. 2021, 44, 113–122. [Google Scholar]
  15. Yeasmin, A.M.; Waliullah, T.M.; Alam, M.A.; Islam, N.; Rahman, A.S. Co-toxicity evaluation of lambda-cyhalothrin and its synergist PBO for susceptibility of Alphitobius diaperinus (Coleoptera: Tenebrionidae). World J. Pharm. Res 2015, 4, 239–253. [Google Scholar]
  16. Li, W.; Naeem, M.; Cui, J.; Du, G.; Chen, H. Toxicity and sublethal effects of lambda-cyhalothrin insecticide on parent and filial generations of Henosepilachna vigintioctomaculata (Coleoptera: Coccinellidae). Insects 2025, 16, 259. [Google Scholar] [CrossRef] [PubMed]
  17. Lambkin, T.A.; Rice, S. Responses of susceptible and cyfluthrin-resistant broiler house populations of lesser mealworm (Coleoptera: Tenebrionidae) to cyhalothrin. J. Econ. Entomol. 2010, 103, 2155–2163. [Google Scholar] [CrossRef] [PubMed]
  18. He, L.M.; Troiano, J.; Wang, A.; Goh, K. Environmental chemistry, ecotoxicity, and fate of lambda-cyhalothrin. Rev. Environ. Contam. Toxicol. 2008, 195, 71–91. [Google Scholar]
  19. Ju, D.; Liu, Y.X.; Liu, X.; Dewer, Y.; Mota-Sanchez, D.; Yang, X.Q. Exposure to lambda-cyhalothrin and abamectin drives sublethal and transgenerational effects on the development and reproduction of Cydia pomonella. Ecotoxicol. Environ. Saf. 2023, 252, 114581. [Google Scholar] [CrossRef]
  20. Kayahan, A. Lethal and sublethal effects of lambda-cyhalothrin on Aphis fabae (Scopoli, 1763), Myzus persicae (Sulzer, 1776) and Acyrthosiphon pisum (Harris, 1776) (Hemiptera: Aphididae). Turk. J. Entomol. 2023, 47, 175–188. [Google Scholar] [CrossRef]
  21. Kavallieratos, N.G.; Boukouvala, M.C.; Eleftheriadou, N.; Xefteri, D.N.; Gidari, D.L.S.; Kyrpislidi, V.P.C. The sublethal impacts of five insecticidal formulations on Oryzaephilus surinamensis behavioral traits. Pest Manag. Sci. 2024, 80, 5334–5341. [Google Scholar] [CrossRef]
  22. Wang, N.; Wang, Z.; Gong, S.; Zhang, Y.; Xue, C. Sublethal concentrations of lambda-cyhalothrin reduce fecundity by affecting the hormone-vitellogenin signaling pathway in Chrysoperla sinica. Entomol. Gen. 2024, 44, 1459–1559. [Google Scholar] [CrossRef]
  23. De França, S.M.; Breda, M.O.; Barbosa, D.R.S.; Araujo, A.M.N.; Guedes, C.A. The sublethal effects of insecticides in insects. In Biological Control of Pest and Vector Insects; Shields, V.D.C., Ed.; BoD–Books on Demand: London, UK, 2017; pp. 23–39. [Google Scholar]
  24. Gidari, D.L.S.; Kavallieratos, N.G.; Boukouvala, M.C. Sublethal effects of α-cypermethrin on the behavioral asymmetries and mating success of Alphitobius diaperinus. Insects 2024, 15, 804. [Google Scholar] [CrossRef]
  25. Campbell, B.; Baldwin, R.; Koehler, P. Locomotion inhibition of Cimex lectularius L. following topical, sublethal dose application of the chitin synthesis inhibitor lufenuron. Insects 2017, 8, 94. [Google Scholar] [CrossRef]
  26. Vallortigara, G.; Chiandetti, C.; Sovrano, V.A. Brain asymmetry (animal). Wiley Interdiscip. Rev. Cogn. Sci. 2011, 2, 146–157. [Google Scholar] [CrossRef]
  27. Anfora, G.; Rigosi, E.; Frasnelli, E.; Ruga, V.; Trona, F.; Vallortigara, G. Lateralization in the invertebrate brain: Left-right asymmetry of olfaction in bumble bee, Bombus terrestris. PLoS ONE 2011, 6, e18903. [Google Scholar] [CrossRef]
  28. Hunt, E.R.; O’Shea-Wheller, T.; Albery, G.F.; Bridger, T.H.; Gumn, M.; Franks, N.R. Ants show a leftward turning bias when exploring unknown nest sites. Biol. Lett. 2014, 10, 20140945. [Google Scholar] [CrossRef] [PubMed]
  29. Güntürkün, O.; Ströckens, F.; Ocklenburg, S. Brain lateralization: A comparative perspective. Physiol. Rev. 2020, 100, 1019–1063. [Google Scholar] [CrossRef]
  30. Rogers, L.J. Brain lateralization and cognitive capacity. Animals 2021, 11, 1996. [Google Scholar] [CrossRef] [PubMed]
  31. Liga, D.; Stancher, G.; Frasnelli, E. Visuo-motor lateralization in Apis mellifera: Flight speed differences in foraging choices. Sci. Rep. 2024, 14, 660. [Google Scholar] [CrossRef]
  32. Romano, D.; Kavallieratos, N.G.; Athanassiou, C.G.; Stefanini, C.; Canale, A.; Benelli, G. Impact of geographical origin and rearing medium on mating success and lateralization in the rice weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae). J. Stored Prod. Res. 2016, 69, 106–112. [Google Scholar] [CrossRef]
  33. Benelli, G.; Romano, D.; Kavallieratos, N.G.; Conte, G.; Stefanini, C.; Mele, M.; Athanassiou, C.G.; Canale, A. Multiple behavioural asymmetries impact male mating success in the khapra beetle, Trogoderma granarium. J. Pest Sci. 2017, 90, 901–909. [Google Scholar] [CrossRef]
  34. Boukouvala, M.C.; Romano, D.; Kavallieratos, N.G.; Stefanini, C.; Canale, A.; Benelli, G. Behavioral asymmetries affecting male mating success in Tenebrio molitor (Coleoptera: Tenebrionidae), an important edible species. J. Econ. Entomol. 2021, 114, 454–461. [Google Scholar] [CrossRef]
  35. Obata, H.; Manabe, A.; Nakamura, N.; Onishi, T.; Senba, Y. A new light on the evolution and propagation of prehistoric grain pests: The World’s oldest maize weevils found in Jomon potteries, Japan. PLoS ONE 2011, 6, e14785. [Google Scholar] [CrossRef]
  36. Cordeiro, E.M.C.; Correa, A.S.; Rosi-Denadai, C.A.; Tome, H.V.V.; Guedes, R.N.C. Insecticide resistance and size assortative mating in females of the maize weevil (Sitophilus zeamais). Pest Manag. Sci. 2017, 73, 823–829. [Google Scholar] [CrossRef]
  37. Arrahman, A.; Mirsam, H.; Djaenuddin, N.; Suriani; Pakki, S.; Saenong, M.S.; Sebayang, A. An in-depth study on Sitophilus zeamais Motsch (Coleoptera: Curculionidae) pests on corn plants. IOP Conf. Ser. Earth Environ. Sci. 2022, 1107, 012060. [Google Scholar] [CrossRef]
  38. Asgher, S.; Avasthi, S. Infestation and growth index on Zea mays by Sitophilus zeamais affected by temperature and humidity; created in in-vitro conditions. Int. J. Entomol. Res. 2024, 9, 74–79. [Google Scholar]
  39. Agathokleous, E.; Blande, J.D.; Calabrese, E.J.; Guedes, R.N.C.; Benelli, G. Stimulation of insect vectors of pathogens by sublethal environmental contaminants: A hidden threat to human and environmental health? Environ. Pollut. 2023, 336, 122422. [Google Scholar] [CrossRef]
  40. Qu, Y.; Xiao, D.; Liu, J.; Chen, Z.; Song, L.; Desneux, N.; Benelli, G.; Xiwu, G.; Song, D. Sublethal and hormesis effects of beta-cypermethrin on the biology, life table parameters and reproductive potential of soybean aphid Aphis glycines. Ecotoxicology 2017, 26, 1002–1009. [Google Scholar] [CrossRef] [PubMed]
  41. Trematerra, P.; Ianiro, R.; Athanassiou, C.G.; Kavallieratos, N.G. Behavioral interactions between Sitophilus zeamais and Tribolium castaneum: The first colonizer matters. J. Pest Sci. 2015, 88, 573–581. [Google Scholar] [CrossRef]
  42. Suleiman, M.; Ibrahim, N.D.; Majeed, Q. Control of Sitophilus zeamais (Motsch) (Coleoptera: Curculionidae) on sorghum using some plant powders. Int. J. Agric. For. 2012, 2, 53–57. [Google Scholar] [CrossRef]
  43. Halstead, D. External sex differences in stored-products Coleoptera. Bull. Entomol. Res. 1963, 54, 119–134. [Google Scholar] [CrossRef]
  44. Tolpo, N.C.; Morrison, E.O. Sex determination by snout characteristics of Sitophilus zeamais Motschulsky. Tex. J. Sci. 1965, 7, 122–124. [Google Scholar]
  45. Boukouvala, M.C.; Romano, D.; Kavallieratos, N.G.; Athanassiou, C.G.; Stefanini, C.; Canale, A.; Benelli, G. Does geographical origin affect lateralization and male mating success in Rhyzopertha dominica beetles? J. Stored Prod. Res. 2020, 88, 101630. [Google Scholar] [CrossRef]
  46. Guedes, N.M.P.; Guedes, R.N.C.; Campbell, J.F.; Throne, J.E. Mating behaviour and reproductive output in insecticide-resistant and-susceptible strains of the maize weevil (Sitophilus zeamais). Ann. Appl. Biol. 2017, 170, 415–424. [Google Scholar] [CrossRef]
  47. Finney, D.J. Statistical Methods in Biological Assay; Charles Griffin: London, UK, 1978. [Google Scholar]
  48. R Core Team. R: A Language and Environment for Statistical Computing; R Foundation for Statistical Computing: Vienna, Austria, 2017; Available online: https://www.Rproject.org/ (accessed on 30 June 2025).
  49. SAS Institute Inc. Using JMP 16.2; SAS Institute Inc.: Cary, NC, USA, 2021. [Google Scholar]
  50. Rogers, L.J.; Vallortigara, G.; Andrew, R.J. Divided Brains: The Biology and Behaviour of Brain Asymmetries, 1st ed.; Cambridge UP: Cambridge, UK, 2013. [Google Scholar]
  51. Boukouvala, M.C.; Kavallieratos, N.G.; Maggi, F.; Angeloni, S.; Ricciutelli, M.; Spinozzi, E.; Ferrati, M.; Petrelli, R.; Canale, A.; Benelli, G. Being exposed to Acmella oleracea-based insecticide extract reduces mobility and mating success in Prostephanus truncatus, the major pest of maize in storages. J. Stored Prod. Res. 2023, 104, 102151. [Google Scholar] [CrossRef]
  52. Walgenbach, C.A.; Burkholder, W.E. Mating behavior of the maize weevil, Sitophilus zeamais (Coleoptera: Curculionidae). Ann. Entomol. Soc. Am. 1987, 80, 578–583. [Google Scholar] [CrossRef]
  53. Kavallieratos, N.G.; Athanassiou, C.G.; Guedes, R.N.; Drempela, J.D.; Boukouvala, M.C. Invader competition with local competitors: Displacement or coexistence among the invasive khapra beetle, Trogoderma granarium Everts (Coleoptera: Dermestidae), and two other major stored-grain beetles? Front. Plant Sci. 2017, 8, 1837. [Google Scholar] [CrossRef]
  54. Hoppe, K.R.; Roush, R.T. Mate finding, dispersal, number released, and the success of biological control introductions. Ecol. Entomol. 1993, 18, 321–331. [Google Scholar] [CrossRef]
  55. Hardy, I.C.; Ode, P.J.; Siva-Jothy, M. Mating behavior. In Insects as Natural Enemies: A Practical Perspective; Jervis, M.A., Ed.; Springer: Dordrecht, The Netherlands, 2005; pp. 219–260. [Google Scholar]
  56. Sallam, M.N. Insect Damage: Damage on Post-Harvest; Food and Agriculture Organization of the United Nations: Rome, Italy, 2013; Volume 38, p. 1. [Google Scholar]
  57. Rosentrater, K.A. Insects in grains: Identification, damage, and detection. In Storage of Cereal Grains and their Products; Rosentrater, K.A., Ed.; Elsevier: Duxford, UK, 2022; pp. 261–292. [Google Scholar]
  58. Ojo, J.A.; Omoloye, A.A. Rearing the maize weevil, Sitophilus zeamais, on an artificial maize cassava diet. J. Insect Sci. 2012, 12, 69. [Google Scholar] [CrossRef]
  59. Chidege, M.Y.; Venkataramana, P.B.; Ndakidemi, P.A. Enhancing food grains storage systems through insect pest detection and control measures for maize and beans: Ensuring food security post-COVID-19 Tanzania. Sustainability 2024, 16, 1767. [Google Scholar] [CrossRef]
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Figure 1. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to control. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 77 pairs).
Figure 1. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to control. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 77 pairs).
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Insects 16 00865 g002
Figure 2. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to λ-cyhalothrin LC10. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 111 pairs).
Figure 2. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to λ-cyhalothrin LC10. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 111 pairs).
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Figure 3. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to λ-cyhalothrin LC30. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 139 pairs).
Figure 3. Flow chart of courtship and mating behavior of Sitophilus zeamais adults exposed to λ-cyhalothrin LC30. Different lateralized traits (i.e., approaching, mounting, and leaving the female’s body) exhibited by males during each behavioral phase are indicated by the color of the boxes: black, green, yellow, and blue for back, left, right, and front side, respectively. Inside each grey box: Y = Yes and N = No (n = 139 pairs).
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Figure 4. Copulation success of Sitophilus zeamais adults exposed to control, or LC10 or LC30 of λ-cyhalothrin. The asterisks indicate significant differences (generalized linear model, binomial distribution, p < 0.01), n.s. = not significant.
Figure 4. Copulation success of Sitophilus zeamais adults exposed to control, or LC10 or LC30 of λ-cyhalothrin. The asterisks indicate significant differences (generalized linear model, binomial distribution, p < 0.01), n.s. = not significant.
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Table 1. Contact toxicity of λ-cyhalothrin on Sitophilus zeamais adults after 72 h of exposure.
Table 1. Contact toxicity of λ-cyhalothrin on Sitophilus zeamais adults after 72 h of exposure.
Active IngredientUnitLC10 (95% CI)LC30 (95% CI)LC50 (95% CI)χ2 (df = 23)p
λ-cyhalothrinmg a.i./cm20.000212 (0.000177–0.000241)0.000282 (0.000250–0.000308)0.000344 (0.000317–0.000366)30.3 1.00
LC = lethal concentration that kills 10%, 30% and 50% of the exposed individuals. 95% CI = lower and upper limits of the 95% confidence interval.
Table 2. Effect of the approaching side on the main mating traits of Sitophilus zeamais exposed to control, or LC10 or LC30 of λ-cyhalothrin.
Table 2. Effect of the approaching side on the main mating traits of Sitophilus zeamais exposed to control, or LC10 or LC30 of λ-cyhalothrin.
Precopula
TraitsDirection of ApproachMate Recognition (min)Interaction (min)Walking (min)Mounting Attempts (Number)Copulation (min)
Treatment
controlBack2.8 ± 0.1 D6.7 ± 0.5 BC7.2 ± 1.2 A1.7 ± 0.2 C268.1 ± 13.6 A
Left1.8 ± 0.1 F3.8 ± 1.0 D4.8 ± 0.3 B1.6 ± 0.1 C276.9 ± 13.9 A
Right2.2 ± 0.1 E6.2 ± 0.6 C5.1 ± 0.4 B1.6 ± 0.1 C274.1 ± 14.2 A
Tested beetles
(number = back + left + right)		11 + 37 + 29 = 7711 + 37 + 29 = 7711 + 37 + 29 = 7711 + 37 + 29 = 777 + 25 + 18 = 50
LC10Back2.8 ± 0.2 CD7.2 ± 0.6 BC3.6 ± 0.3 D4.2 ± 0.8 B109.2 ± 28.0 C
Left2.7 ± 0.1 D5.9 ± 0.4 C4.2 ± 0.2 C3.9 ± 0.5 B160.4 ± 19.2 B
Right3.0 ± 0.1 C6.5 ± 0.5 C3.7 ± 0.2 D3.9 ± 0.5 B82.4 ± 21.4 C
Front3.4 ± 0.3 BC8.9 ± 1.2 AB4.1 ± 0.4 BCD5.8 ± 1.2 AB46.8 ± 6.7 D
Tested beetles
(number = back+ left + right + front)		18 + 47 + 39 + 7 = 11118 + 47 + 39 + 7 = 11118 + 47 + 39 + 7 = 11118 + 47 + 39 + 7 = 1116 + 27 + 10 + 7= 50
LC30Back3.9 ± 0.2 A8.5 ± 0.6 AB3.1 ± 0.2 D4.8 ± 0.6 B95.1 ± 24.2 C
Left3.5 ± 0.1 B6.2 ± 0.4 C3.7 ± 0.2 D4.6 ± 0.5 B119.1 ± 17.4 C
Right3.5 ± 0.1 B7.6 ± 0.5 B3.3 ± 0.1 D4.1 ± 0.5 B80.4 ± 19.1 C
Front3.7 ± 0.2 B9.7 ± 0.7 A3.32 ± 0.3 D6.5 ± 0.6 A18.6 ± 18.6 E
Tested beetles
(number = back+ left + right + front)		21 + 58 + 46 + 14 = 13921 + 58 + 46 + 14 = 13921 + 58 + 46 + 14 = 13921 + 58 + 46 + 14 = 1394 + 27 + 18 + 1 = 50
χ2, df, p170.0, 10, <0.0181.6, 10, <0.0163.7, 10, <0.0196.7, 10, <0.01140.5, 10, <0.01
Values are means (±standard errors). Per trait, within each column, different letters indicate significant differences (Steel–Dwass test, p < 0.05).
Table 3. Effect of the mounting side on the main mating traits of Sitophilus zeamais exposed to control, or LC10, or LC30 of λ-cyhalothrin.
Table 3. Effect of the mounting side on the main mating traits of Sitophilus zeamais exposed to control, or LC10, or LC30 of λ-cyhalothrin.
Precopula
TraitsDirection of Mounting SideMate Recognition (min)Interaction (min)Walking (min)Mounting Attempts
(Number)		Copulation (min)
Treatment
controlBack2.1 ± 0.2 E5.4 ± 0.7 C5.3 ± 0.6 A1.7 ± 0.2 D248.4 ± 10.8 B
Left1.9 ± 0.1 E4.5 ± 0.3 D5.3 ± 0.4 A1.7 ± 0.1 D269.0 ± 10.1 B
Right2.0 ± 0.2 E6.2 ± 0.8 BC5.2 ± 0.5 A1.6 ± 0.1 D309.7 ± 24.1 A
Tested beetles
(number = back + left + right)		11 + 37 + 29 = 7711 + 37 + 29 = 7711 + 37 + 29 = 7711 + 37 + 29 = 777 + 25 + 18 = 50
LC10Back2.8 ± 0.2 CD7.2 ± 0.6 BC3.8 ± 0.3 BC4.4 ± 1.0 B115.5 ± 8.2 D
Left2.7 ± 0.1 D5.9 ± 0.4 C4.1 ± 0.3 B3.6 ± 0.5 C133.9 ± 5.3 C
Right3.0 ± 0.1 C6.5 ± 0.5 BC3.6 ± 0.2 BC4.1 ± 0.6 C103.2 ± 5.0 D
Front3.4 ± 0.3 ABC8.9 ± 1.2 AB4.1 ± 0.4 B5.3 ± 1.1 AB61.8 ± 14.9 E
Tested beetles
(number = back+ left + right + front)		20 + 47 + 36 + 8 = 11120 + 47 + 36 + 8 = 11120 + 47 + 36 + 8 = 11120 + 47 + 36 + 8 = 1119 + 29 + 11 + 1 = 50
LC30Back3.9 ± 0.2 A7.8 ± 0.5 B3.3 ± 0.2 C4.8 ± 0.6 B104.3 ± 6.8 D
Left3.5 ± 0.1 B6.2 ± 0.4 C3.6 ± 0.2 BC4.6 ± 0.5 B121.0 ± 2.0 D
Right3.5 ± 0.1 B7.6 ± 0.6 B3.3 ± 0.1 C4.1 ± 0.5 BC51.8 ± 6.7 E
Front3.7 ± 0.2 AB9.9 ± 0.7 A3.3 ± 0.3 C6.5 ± 0.6 A44.6 ± 26.0 E
Tested beetles
(number = back+ left + right + front)		26 + 56 + 42 + 15 = 13926 + 56 + 42 + 15 = 13926 + 56 + 42 + 15 = 13926 + 56 + 42 + 15 = 13910 + 26 + 13 + 1 = 50
χ2, df, p170.5, 10, <0.0159.5, 10, <0.0159.4, 10, <0.01126.8, 10, <0.01129.6, 10, <0.01
Values are means (±standard errors). Per trait, within each column, different letters indicate significant differences (Steel-Dwass test, p < 0.05).
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Boukouvala, M.C.; Kavallieratos, N.G.; Gidari, D.L.S.; Filintas, C.S.; Skourti, A.; Kyrpislidi, V.P.C.; Skordos, D.P. Love in the Time of Pyrethroids: Mating Behavior of Sitophilus zeamais Is Influenced by Sublethal Concentrations of λ-Cyhalothrin and Lateralization. Insects 2025, 16, 865. https://doi.org/10.3390/insects16080865

AMA Style

Boukouvala MC, Kavallieratos NG, Gidari DLS, Filintas CS, Skourti A, Kyrpislidi VPC, Skordos DP. Love in the Time of Pyrethroids: Mating Behavior of Sitophilus zeamais Is Influenced by Sublethal Concentrations of λ-Cyhalothrin and Lateralization. Insects. 2025; 16(8):865. https://doi.org/10.3390/insects16080865

Chicago/Turabian Style

Boukouvala, Maria C., Nickolas G. Kavallieratos, Demeter Lorentha S. Gidari, Constantin S. Filintas, Anna Skourti, Vasiliki Panagiota C. Kyrpislidi, and Dionysios P. Skordos. 2025. "Love in the Time of Pyrethroids: Mating Behavior of Sitophilus zeamais Is Influenced by Sublethal Concentrations of λ-Cyhalothrin and Lateralization" Insects 16, no. 8: 865. https://doi.org/10.3390/insects16080865

APA Style

Boukouvala, M. C., Kavallieratos, N. G., Gidari, D. L. S., Filintas, C. S., Skourti, A., Kyrpislidi, V. P. C., & Skordos, D. P. (2025). Love in the Time of Pyrethroids: Mating Behavior of Sitophilus zeamais Is Influenced by Sublethal Concentrations of λ-Cyhalothrin and Lateralization. Insects, 16(8), 865. https://doi.org/10.3390/insects16080865

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