Tritrophic effects of different cereal flour on life table parameters and population projection of Habrobracon hebetor (Hymenoptera: Braconidae)

Tritrophic effects of different cereal flour on life table parameters and population projection of Habrobracon hebetor (Hymenoptera: Braconidae) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Tritrophic effects of different cereal flour on life table parameters and population projection of Habrobracon hebetor (Hymenoptera: Braconidae) Narges Mostafazadeh, Fariba Mehrkhou, Nabat Naghshbandi, Remzi Atlihan This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4632393/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 The parasitoid wasp, Habracon hebetor , Say is used widely in mass production against larval stage of field and stored product insect pests. Less is known about the third-trophic level effects of cereal flour on the population dynamics of H. hebetor . In this study, tritrophic level effects of three cereal flours (wheat, corn, and barley) on population dynamics and population projection of H. hebetor were studied. Data were analyzed by constructing the age-stage, two-sex life table, to take into consideration either female or male and stage differentiation. According to our results, the used corn host type ( var .704) was more suitable host for H. hebetor than wheat and barley due to the shortening developmental time (10.76 ± 0.08 d), increasing survival rate (0.74 ± 0.006), female longevity (14.93 ± 0.92 d) and fecundity (91.20 ± 9.05 offspring/female). Also, the population growth parameters, including the intrinsic rate of increase (0.2217 ± 0.0178 d − 1 ), finite rate of increase (1.25 ± 0.02 d − 1 ), and reproductive rate (27.92 ± 6.57 offspring) were the highest on corn host type. Interestingly, the predicted of population growth of H. hebetor was the fastest and highest on corn, followed by wheat, and barley. The results of this study provide direction to design a comprehensive program for the mass rearing of H. hebetor . Biological sciences/Biological techniques Biological sciences/Developmental biology Biological sciences/Ecology Ectoparasitoid wasp Cereal flour Demographic parameters Age-stage two-sex life table Population projection Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Habrobracon hebetor Say (Hymenoptera: Braconidae) as an ectoparasitoid, is used widely in host-parasitoid interaction studies due to the high reproduction rate, short generation period, and wide range of hosts, as well as its high aggressiveness abilities 1 . Additionally, in biological pest management programs, it is considered along with the release of egg and young larval parasitoids 2 . It has been used in biological control against larvae of field and storage lepidopteran pests, including Tuta absoluta Meyrick, Spodoptera litura Fabricius, Helicoverpa armigera Hübner 3–7 , Ephestia kuehniella Zeller, Plodia interpunctella Hübner Galleria mellonella L. and Sitotroga cerealella Olivier 8–10 . Recently, there has been an increasing interest in the third-trophic level effects among host plants, phytophagous pests, and their natural enemies 7, 11–12 . In most studies, the importance of the host plant on different properties of phytophagous pests has been considered. For example, the effects of various cereal flour on the life table parameters of E. kuehniella showed that corn was the most suitable host for the Mediterranean flour moth 13 . Also, 14 stated that there was a relationship between the quality of the host plant and the growth indices of E.kuehniella , which is fed on corn (var. 704). In some studies, the suitability of the second trophic level (i.e., host species) on the biological, physiological, and behavioral aspects of H. hebetor was discussed. For example, the effects of stored insect species, including E. kuehniella and G. mellonella larvae, on the demographic parameters of H. hebetor were studied by 4 . Saadat et al. 6 reported the developmental time and digestive enzymes activity of H. hebetor which reared on Ectomyelois ceratoniae Zeller, P. interpunctella , E. kuehniella , H. armigera , and Malacosoma disstria Hübner. They found that, the stored product insects were more suitable for H. hebetor , due to the highest content of sugar and glycogen in stored insect pests than the field crop insects. Despite the importance of host type, less is known about the third-trophic level effects on the biological, population, and projection characteristics of H. hebetor . However, the effects of four host types on the biological properties of H. hebetor were discussed by 15 . Also, our previous study showed the importance of host plant type on the behavioral properties, including the instantaneous attack rate and the handling time of H. hebetor , who showed that corn (var.704) was more effective host plant type on the parasitoid’s functional response parameters 16 . Along with our previous studies, we have received attention to demonstrate the third-trophic level effects of the same used host plants on the population growth parameters and population projection using age-stage, two-sex life table. By constructing this theory, not only the male population did not ignored, but also the stage differentiation can be considered 17 . Amir-Maafi & Chi 4 constructed age-stage, two-sex life table to evaluate the effects of E. kuehniella and G. mellonella on the population properties of H. hebetor , which fed on wheat flour. According to their reports, the aforementioned lepidopteran insects did not affect the developmental time and population growth parameters of H. hebetor . To our knowledge, there are no documents regarding the third-trophic level effects of cereal flour on the life table parameters of H. hebetor . Demographic information is not only helpful for designing the mass production programs of H. hebetor , but also the obtained data give more comprehensive information regarding projection of population growth 4 . Results Biological parameters . The third-trophic level effects on the biological parameters (e.g., developmental time, survival, adult longevity, reproduction, and total life span) of H. hebetor are shown in Table 1. Feeding of E. kuehniella on different cereal flour did not change the egg and larval duration of the parasitoid. However, the pupal, preadult period and, survival of preadult stages of H.hebetor were affected by the type of cereal flour. The highest pupa and preadult duration of H.hebetor were obtained on wheat. Also, female longevity was affected by the cereal flour, in which the most extended female longevity of parasitoids was obtained on corn, followed by barley and wheat. Male longevity was shorter than females in all treatments. There were significant differences in reproductive days and reproduction rate, which were the highest on the corn host type. The third-trophic level effects of three cereal flour on the stage differentiation and survivorship curves of the F 1 generation of H. hebetor are depicted in Fig. 1 . The age-stage specific survival rate ( s xj ) not only shows the probability achivement of an individual egg of parasitoid for surviving in age x and stage j , but also it gives a precise description about the survival and stage transitions, and the stage overlap during the development of a cohort, as well. The reproduction of parasitoids began at age 9 d in all treatments. The age-specific fecundity ( m x = 7.07 eggs) and age-specific maternity ( l x m x = 3.95 offspring) peaks of the females were higher on corn than those of observed for other progeny groups of cereal flour adults. The lowest net maternity peak ( l x m x = 3.24 offspring) was obtained for females on barley at age 11 d (Fig. 2 ). The age stage specific life expectancy ( e xj ) shows the third-trophic level effects of cereal flour on the predicted lifespan of H.hebetor individual. The life expectancy of a parasitoids’ neonate was 13.57, 15.82, and 12.20 d on wheat, corn, and barley, respectively (Fig. 3 ). The used cereal flour affected the age stage-specific reproductive value ( v xj ) of H. hebetor (Fig. 4 ). The earliest and highest reproductive value peaks were observed on barley ( v 10 = 44.60). The lowest peak of reproductive value was obtained on corn ( v 12 = 8.05). Population growth parameters and Population projection. The third-trophic level effects of used cereal flour on the population parameters of H.hebetor using a paired bootstrap test (at 5% level) are demonstrated in Table 2. The highest values of the net reproductive rate ( R 0 ), the intrinsic rate of increase ( r ), and the finite rate of increase (λ) were observed on corn. The effects of cereal flour on the projected population growth of H. hebetor are shown in Fig. 5 . The highest population size was on the corn, then followed by wheat and barley during 60 days (Fig. 5 ). Also, the range of variability host type in population growth indicated that the lowest degree of uncertainty in the population growth of the parasitoid was on corn (Fig. 6 ). Discussion Biological and population traits of parasitoids depend on several factors such as, the host plant, host species and, it’s physiological condition 7,11, 18–20 . The type of host plants involved in the developmental time period of parasitoids by shortening their preadults duration, increasing survival rate, adult body size, adult longevity, reproduction days and, fecundity 21 as well as increasing the population growth parameters 4 . This research was studied owing to the importance of host plant type on the foraging behavior, physiological and population dynamics of parasitoids 6, 22, 23 , and consequently to have successful production of biocontrol agents. Alongside of our previous study 16 , the current research was reported the third-trophic level effects of the cereal flour on the biological, life table traits, and population projection of H. hebetor . Our results demonstrated that usage of corn and barley cereal flour, accelerated the immature developmental time of H. hebetor , and increased the survival rate, adult longevity as well as reproduction days. Amir-Maafi & Chi 4 reported a developmental time of 10.83 d for H. hebetor on the fifth instar larvae of E. kuehniella fed on wheat. The preadult periods of the parasitoid in our study are within the same range as this report. However, 7 obtained a developmental time of 17.05 d of H. hebetor on the cotton bollworm, H. armigera , which fed on the leaf and fruit of tomato, it wasmore than in our study. Saadat et al. 6 concluded that host plants are rich in sugar and glycogen and provide more suitable physiological conditions than host species that contain terpenes and tannins for H. hebetor. Our results showed that the total number of eggs/female during the lifetime of H. hebetor varied from 70.87 to 91.20 eggs/female on E. kuehniella , which values are much higher than the 66.30 eggs/female on E.kuehniella 4 and 61.62 eggs/female on H. armigera 7 . Variation in preadult developmental time and fecundity rates among these studies could be because of different host species, quality of host type, rearing conditions, and physiological condition of host species 4,6,14 . Seyedi et al. 14 stated that the highest protein content of corn (var. 704) resulted in the highest protein content and proteolytic activity of E. kuehniella . Similarly, Saadat et al. 6 stated that the quality and quantity of protease and amylase activity of H. hebector guts was greatest when reared on E. kuehniella . This fact implies the importance of the quality and quantity of host type on the physiological condition at the tritrophic level among host plants, phytophagous pests, and their natural enemies 11,12 . Construction and analyzing the life table data based on the age-stage, two-sex life table 24,25 , not only consider the importance of each stages on the population dynamics,, but also it did not ignore the male population 17,26 . Our results demonstrated that the corn variety used in this research, was found to be more effective on H. hebetor population growth rate of offspring by increasing the net reproductive rate ( R 0 ), the intrinsic rate of increase ( r ), the finite rate of increase ( λ ), and shortening the mean generation time ( T ). The highest rate of intrinsic rates of increase on corn reflected the combined effects of age-specific fecundity ( m x ) and age-specific survival rate ( l x ) 27 . The population growth parameters of the parasitoid in our study are more than those of Amir-Maafi & Chi 4 study. These variations might be due to the variety of host types. The uncertainty based on the 2.5th and 97.5th percentiles of the net reproductive rate ( R 0 ) is another advantage of the age-stage, two-sex life table theory. Our results showed the same trends of population projection with population parameters. The predicted population of H. hebetor was the fastest and highest on corn, followed by wheat, and barley, which was reported for the first time in this study. In conclusion, corn is a more suitable host than wheat and barley for the parasitoid, which is reared on E. kuehniella . It is concluded that corn type, by providing more suitable nutrition, resulted in a shortening the developmental time, increasing the fecundity and survival rate. Interestingly, the suitability of the corn variety in this study, was confirmed once more by the estimation of population projection. Material and Methods Cereal flour and insects rearing. The current study was designed along with our previous studies 16 . In this study, we used the same cereal flour including, wheat ( var . Homma), barley ( var . Makui) and, corn (var. 704), to assign how the host type affects the efficacy of parasitoid, H. hebector . Aforementioned seeds were powdered using grinder and kept in a refrigerator (4 ºC) to avoid any contamination 28 . According to Tarlack et al. 28 method, the rearing of Mediterranean flour moth has been conducted on the aforementioned different cereal flour. For parasitoid rearing, the initial colony of H. hebetor was prepared kindly from the insectarium of Agriculture Research Center, West Azerbaijan, Iran. To obtain the same aged of parasitoid’s egg, the obtained adults of H. hebetor were released on the fifth instar larvae of E.kuehniella , as preferred instar larvae 4 for 24 h. The insects rearing were continued for two generations before using in experiments at controlled condition (25 ± 1ºC, 60 ± 5% R. H., and a photoperiod of 14: 10 h. (L: D)). Developmental time and life table studies. To study biology and demographic parameters of H. hebetor , one egg of parasitoid was left on each larva of E. kuehniella and, the rest of eggs were removed 4 . Hence, the current study was used 49, 49 and, 58 numbers of the fifth instar parasitized larvae of E. kuehniella which obtained on wheat, corn and, barley, respectively. Parasitized larvae containing one egg of H. hebetor were transferred individually to plastic Petri dishes (3 cm in height, 8 cm in diameter). The developmental stage of each individual was recorded daily. By emerging adults, they were paired in plastic containers (4 cm in diameter, 10 cm in height) and fed by 10% honey solution during the oviposition period. According to our previous study 16 , four numbers of fifth instar larvae of E. kuehniella were supplied daily to parasitize during the oviposition period. The survival, adult longevity, oviposition period and, fecundity were recorded daily. All experiments were conducted in laboratory conditions as mentioned earlier. Data analysis. The third-trophic level effects of three different cereal flour on the developmental time, survival, and reproduction of H. hebetor were analyzed according to the age-stage, two-sex life table theory 24,29 using the computer program TWOSEX-MSChart 30 . The bootstrap technique was used to estimate variances and standard errors of life history and population parameters 31,32 . The differences between treatments were assessed using the paired bootstrap test based on the confidence interval of differences 33,34 . The population growth of the parasitoid cohorts was projected according to the technique developed by Chi 35 using the computer program TIMING-MSChart 36 . The life tables based on the 0.025th and 0.975th bootstrap results of the net reproductive rate ( R 0 ), were used to project the uncertainty of the population growth of H.hebetor 37 . Declarations Conflict of Interest There is no conflict of interest among authors. Funding This project was funded by Urmia University Author Contribution N. M.and N. N. conducted the experiments. F. M.designed the experiments, analysed data, and wrote the manuscript. R. A. edited the manuscript. Acknowledgement We are appreciated to Urmia University for financial support. Data Availability All data generated or analysed during this study are included in this published article References Gunduz, E. A. & Gulel, A. Effects of adult age and species on development period of parasitoid Bracon hebetor Say (Hymenoptera: Braconidae). J. Agri. Sci, 20, 31–36 (2005). Tuerxun, A., Guo, W. C., Akedan, L. H. B., Xu, J. J. & He, J. Description of frequent species of Braconid in cotton field in Xinjiang China. Xinjiang Agri. sci, 43 (6), 503–506 (2006). Hopper, K. R. United States Department of Agriculture Agricultural research service research on biological control of arthropods. Pest Manag. Sci, 59, 643–653 (2003). Amir-Maafi, M. & Chi, H. 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Third-trophic level effects (Mean± SE) of cereal flour on the life history parameters of Habrobracon hebetor reared on 5 th instar larvae of Ephestia kuehniella Treatments (n * ) Life history parameters Wheat Corn Barley Egg 2.00±0.00 a (37) 2.00±0.00 a (46) 2.00±0.00 a (38) Larvae 1.83±0.12 a (29) 1.97±0.14 a (36) 1.88±0.13 a (32) Pupa 7.66±0.16 a (29) 6.80±0.17 b (34) 6.84±0.16 b (32) Pre-adult 11.48±0.10 a (29) 10.76±0.08 b (34) 10.72±0.09 c (32) Pre-adult survival rate 0.78±0.007 b (29) 0.74±0.006 a (34) 0.84±0.006 a (32) Female longevity Male longevity Female total longevity Male total longevity 11.00±1.03 b (15) 7.57±0.81 a (14) 22.20±1.09 b (15) 19.36±0.80 a (14) 14.93±0.92 a (15) 5.88±0.34 b (19) 25.93±0.95 a (15) 16.47±0.34 b (19) 13.06±1.10 ab (15) 5.89±0.30 ab (17) 23.80±1.15 ab (15) 16.30±0.32 b (17) APRP 0.33±0.13 a (15) 0.33±0.13 a (15) 0.20±0.11 a (15) TPRP 11.53±0.17 a (15) 11.33±0.20 a (15) 10.93±0.18 b (15) Reproductive days 8.13±0.83 ab (15) 10.60±0.70 a (15) 8.93±0.52 a (15) Reproduction (offspring/female) 70.87±8.23 b (15) 91.20±9.05 a (15) 79.73±7.27 ab (15) Means in the same row followed by different letter are significantly different (paired bootstrap test at 5% significant level). Standard errors were measured by 100,000 bootstrap resampling. APRP=Adult pre-reproductive day; TPRP=Total pre-reproductive day; *n=sample size of each stage of Habrobracon hebetor on each treatment Table 2. Third-trophic level effects (Mean± SE) of cereal flour on the population growth parameters of Habrobracon hebetor reared on 5 th instar larvae of Ephestia kuehniella . Population growth parameters Wheat Treatments (n * ) Corn Barley R 0 (offspring/individual) 21.70±5.26 b (49) 27.92±6.57 a (49) 20.62±4.94 b (58) r (day -1 ) 0.2030±0.0171 b (49) 0.2217±0.0178 a (49) 0.2063±0.0178 b (58) λ (day -1 ) 1.23±0.02 b (49) 1.25±0.02 a (49) 1.23±0.02 b (58) T (day) 15.16±0.38 a (49) 15.02±0.24 b (49) 14.66±0.22 b (58) The means followed by different letters in each raw are significantly different (paired- bootstrap at 5% significance level). Standard errors were measured by 100,000 bootstrap resampling. R 0 = net reproductive rate; r = intrinsic rate of increase; λ = finite rate of increase; T = mean generation time; * n = sample size of each stage of Habrobracon hebetor on each treatment Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-4632393","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":329476135,"identity":"ca5763c5-5a39-4c3f-86c9-5f589362b10c","order_by":0,"name":"Narges Mostafazadeh","email":"","orcid":"","institution":"Urmia University","correspondingAuthor":false,"prefix":"","firstName":"Narges","middleName":"","lastName":"Mostafazadeh","suffix":""},{"id":329476136,"identity":"01d0a1f3-7006-42ca-9aaa-f85f95474f5d","order_by":1,"name":"Fariba Mehrkhou","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA40lEQVRIiWNgGAWjYBAC9gYGBmYGhoQEIHUAyJeACCfg0cJzAK6FLYFkLTwGxDmMh4E78XHhnrQ8fomcb9KVORbyDOyHHzA83INPC+9m4xnPcoolZ+Rukzy7TcKwgSfNgCHhGW4t9gy826R5DlQkbrgB1NK4TYKxgSEH6NADeG3Z/huiJecZSIt9A/8bglq2MfMcyAFpYQNpSWyQIGQLM+9moMPSEmf2PDO2BGpJbpN4ZnAArxb23o2feQ4kJ/azJz+82bitzrafP/nhwx94tIAiBRWwATE+DaNgFIyCUTAKiAAAEadNdY4T208AAAAASUVORK5CYII=","orcid":"","institution":"Urmia University","correspondingAuthor":true,"prefix":"","firstName":"Fariba","middleName":"","lastName":"Mehrkhou","suffix":""},{"id":329476137,"identity":"d2a79249-e06f-4d38-9b18-06a0a6e9de9e","order_by":2,"name":"Nabat Naghshbandi","email":"","orcid":"","institution":"Oshnavieh University","correspondingAuthor":false,"prefix":"","firstName":"Nabat","middleName":"","lastName":"Naghshbandi","suffix":""},{"id":329476138,"identity":"0e0a0ae2-0978-49ef-b585-46aa72ca6923","order_by":3,"name":"Remzi Atlihan","email":"","orcid":"","institution":"University of Van Yuzuncu Yil","correspondingAuthor":false,"prefix":"","firstName":"Remzi","middleName":"","lastName":"Atlihan","suffix":""}],"badges":[],"createdAt":"2024-06-24 22:08:21","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4632393/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4632393/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":60932123,"identity":"82f8788e-0f4d-4ebf-81f1-ebe8cca68efc","added_by":"auto","created_at":"2024-07-23 17:22:59","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":161170,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the age‐stage‐specific survival rate (\u003cem\u003es\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eHabrobracon hebetor\u003c/em\u003e on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of\u003cem\u003e Ephestia kuehniella.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/0c067ff0548051b9c8194e0e.jpeg"},{"id":60932127,"identity":"665e46d8-2329-430e-9b9f-83775b09a466","added_by":"auto","created_at":"2024-07-23 17:22:59","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":150626,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the age‐specific survival rate (\u003cem\u003el\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e), fecundity (\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e), and net maternity (\u003cem\u003el\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eHabrobracon hebetor\u003c/em\u003e on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of\u003cem\u003e Ephestia kuehniella.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/435097c115712ad83207c457.jpeg"},{"id":60932889,"identity":"5e748a05-1ed9-4447-b385-2645be98eda6","added_by":"auto","created_at":"2024-07-23 17:38:59","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":144318,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the age-stage- specific life expectancy (\u003cem\u003ee\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eHabrobracon hebetor \u003c/em\u003ereared on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of\u003cem\u003e Ephestia kuehniella.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/bc1d7b2965a5a4fb014e08cd.jpeg"},{"id":60932559,"identity":"dc87984d-a740-4903-9b17-44479d54f9aa","added_by":"auto","created_at":"2024-07-23 17:30:59","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":103656,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the age-stage-specific reproductive value (\u003cem\u003ev\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eHabrobracon hebetor \u003c/em\u003ereared on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of\u003cem\u003e Ephestia kuehniella.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/5980f1f71f3d4c8ae6ba5d8d.jpeg"},{"id":60932124,"identity":"44424dff-f32e-4e78-a1fa-ace825bdfab2","added_by":"auto","created_at":"2024-07-23 17:22:59","extension":"jpeg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":162856,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the projection of population growth potential and stage structure of \u003cem\u003eHabrobracon hebetor \u003c/em\u003ereared on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of \u003cem\u003eEphestia kuehniella \u003c/em\u003eduring 60 days.\u003c/p\u003e","description":"","filename":"floatimage5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/8e8ee9218a64380671226288.jpeg"},{"id":60932128,"identity":"c76a1a6e-df55-4406-9fbd-039584604da0","added_by":"auto","created_at":"2024-07-23 17:22:59","extension":"jpeg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":143317,"visible":true,"origin":"","legend":"\u003cp\u003eThird-trophic level effects of the cereal flour on the confidence interval of the population growth of \u003cem\u003eHabrobracon hebetor \u003c/em\u003ereared on the 5\u003csup\u003eth\u003c/sup\u003e instar larvae of\u003cem\u003e Ephestia kuehniella\u003c/em\u003e during 60 days, the cohorts were constructed based on the 2.5 and 97.5% percentiles of net reproductive rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e).\u003c/p\u003e","description":"","filename":"floatimage6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/fed803ec7f5d5b9bc4e84ae8.jpeg"},{"id":64309300,"identity":"dd078ef1-9c7c-4687-bd52-d66c56b9a672","added_by":"auto","created_at":"2024-09-11 13:14:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1432764,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4632393/v1/3fb5389a-0e8f-4517-8b0d-3e406e849406.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Tritrophic effects of different cereal flour on life table parameters and population projection of Habrobracon hebetor (Hymenoptera: Braconidae)","fulltext":[{"header":"Introduction","content":"\u003cp\u003e \u003cem\u003eHabrobracon hebetor\u003c/em\u003e Say (Hymenoptera: Braconidae) as an ectoparasitoid, is used widely in host-parasitoid interaction studies due to the high reproduction rate, short generation period, and wide range of hosts, as well as its high aggressiveness abilities\u003csup\u003e1\u003c/sup\u003e. Additionally, in biological pest management programs, it is considered along with the release of egg and young larval parasitoids\u003csup\u003e2\u003c/sup\u003e. It has been used in biological control against larvae of field and storage lepidopteran pests, including \u003cem\u003eTuta absoluta\u003c/em\u003e Meyrick, \u003cem\u003eSpodoptera litura\u003c/em\u003e Fabricius, \u003cem\u003eHelicoverpa armigera\u003c/em\u003e H\u0026uuml;bner\u003csup\u003e3\u0026ndash;7\u003c/sup\u003e, \u003cem\u003eEphestia kuehniella\u003c/em\u003e Zeller, \u003cem\u003ePlodia interpunctella\u003c/em\u003e H\u0026uuml;bner \u003cem\u003eGalleria mellonella\u003c/em\u003e L. and \u003cem\u003eSitotroga cerealella\u003c/em\u003e Olivier\u003csup\u003e8\u0026ndash;10\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eRecently, there has been an increasing interest in the third-trophic level effects among host plants, phytophagous pests, and their natural enemies\u003csup\u003e7, 11\u0026ndash;12\u003c/sup\u003e. In most studies, the importance of the host plant on different properties of phytophagous pests has been considered. For example, the effects of various cereal flour on the life table parameters of \u003cem\u003eE. kuehniella\u003c/em\u003e showed that corn was the most suitable host for the Mediterranean flour moth\u003csup\u003e13\u003c/sup\u003e. Also, \u003csup\u003e14\u003c/sup\u003e stated that there was a relationship between the quality of the host plant and the growth indices of \u003cem\u003eE.kuehniella\u003c/em\u003e, which is fed on corn (var. 704). In some studies, the suitability of the second trophic level (i.e., host species) on the biological, physiological, and behavioral aspects of \u003cem\u003eH. hebetor\u003c/em\u003e was discussed. For example, the effects of stored insect species, including \u003cem\u003eE. kuehniella\u003c/em\u003e and \u003cem\u003eG. mellonella\u003c/em\u003e larvae, on the demographic parameters of \u003cem\u003eH. hebetor\u003c/em\u003e were studied by\u003csup\u003e4\u003c/sup\u003e. Saadat et al. \u003csup\u003e6\u003c/sup\u003e reported the developmental time and digestive enzymes activity of \u003cem\u003eH. hebetor\u003c/em\u003e which reared on \u003cem\u003eEctomyelois ceratoniae\u003c/em\u003e Zeller, \u003cem\u003eP. interpunctella\u003c/em\u003e, \u003cem\u003eE. kuehniella\u003c/em\u003e, \u003cem\u003eH. armigera\u003c/em\u003e, and \u003cem\u003eMalacosoma disstria\u003c/em\u003e H\u0026uuml;bner. They found that, the stored product insects were more suitable for \u003cem\u003eH. hebetor\u003c/em\u003e, due to the highest content of sugar and glycogen in stored insect pests than the field crop insects.\u003c/p\u003e \u003cp\u003eDespite the importance of host type, less is known about the third-trophic level effects on the biological, population, and projection characteristics of \u003cem\u003eH. hebetor\u003c/em\u003e. However, the effects of four host types on the biological properties of \u003cem\u003eH. hebetor\u003c/em\u003e were discussed by \u003csup\u003e15\u003c/sup\u003e. Also, our previous study showed the importance of host plant type on the behavioral properties, including the instantaneous attack rate and the handling time of \u003cem\u003eH. hebetor\u003c/em\u003e, who showed that corn (var.704) was more effective host plant type on the parasitoid\u0026rsquo;s functional response parameters\u003csup\u003e16\u003c/sup\u003e. Along with our previous studies, we have received attention to demonstrate the third-trophic level effects of the same used host plants on the population growth parameters and population projection using age-stage, two-sex life table. By constructing this theory, not only the male population did not ignored, but also the stage differentiation can be considered\u003csup\u003e17\u003c/sup\u003e. Amir-Maafi \u0026amp; Chi \u003csup\u003e4\u003c/sup\u003e constructed age-stage, two-sex life table to evaluate the effects of \u003cem\u003eE. kuehniella\u003c/em\u003e and \u003cem\u003eG. mellonella\u003c/em\u003e on the population properties of \u003cem\u003eH. hebetor\u003c/em\u003e, which fed on wheat flour. According to their reports, the aforementioned lepidopteran insects did not affect the developmental time and population growth parameters of \u003cem\u003eH. hebetor\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eTo our knowledge, there are no documents regarding the third-trophic level effects of cereal flour on the life table parameters of \u003cem\u003eH. hebetor\u003c/em\u003e. Demographic information is not only helpful for designing the mass production programs of \u003cem\u003eH. hebetor\u003c/em\u003e, but also the obtained data give more comprehensive information regarding projection of population growth\u003csup\u003e4\u003c/sup\u003e.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e \u003cb\u003eBiological parameters\u003c/b\u003e. The third-trophic level effects on the biological parameters (e.g., developmental time, survival, adult longevity, reproduction, and total life span) of \u003cem\u003eH. hebetor\u003c/em\u003e are shown in Table\u0026nbsp;1. Feeding of \u003cem\u003eE. kuehniella\u003c/em\u003e on different cereal flour did not change the egg and larval duration of the parasitoid. However, the pupal, preadult period and, survival of preadult stages of \u003cem\u003eH.hebetor\u003c/em\u003e were affected by the type of cereal flour. The highest pupa and preadult duration of \u003cem\u003eH.hebetor\u003c/em\u003e were obtained on wheat. Also, female longevity was affected by the cereal flour, in which the most extended female longevity of parasitoids was obtained on corn, followed by barley and wheat. Male longevity was shorter than females in all treatments. There were significant differences in reproductive days and reproduction rate, which were the highest on the corn host type.\u003c/p\u003e \u003cp\u003eThe third-trophic level effects of three cereal flour on the stage differentiation and survivorship curves of the F\u003csub\u003e1\u003c/sub\u003e generation of \u003cem\u003eH. hebetor\u003c/em\u003e are depicted in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. The age-stage specific survival rate (\u003cem\u003es\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) not only shows the probability achivement of an individual egg of parasitoid for surviving in age \u003cem\u003ex\u003c/em\u003e and stage \u003cem\u003ej\u003c/em\u003e, but also it gives a precise description about the survival and stage transitions, and the stage overlap during the development of a cohort, as well. The reproduction of parasitoids began at age 9 d in all treatments. The age-specific fecundity (\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e = 7.07 eggs) and age-specific maternity (\u003cem\u003el\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e = 3.95 offspring) peaks of the females were higher on corn than those of observed for other progeny groups of cereal flour adults. The lowest net maternity peak (\u003cem\u003el\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e = 3.24 offspring) was obtained for females on barley at age 11 d (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The age stage specific life expectancy (\u003cem\u003ee\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) shows the third-trophic level effects of cereal flour on the predicted lifespan of \u003cem\u003eH.hebetor\u003c/em\u003e individual. The life expectancy of a parasitoids\u0026rsquo; neonate was 13.57, 15.82, and 12.20 d on wheat, corn, and barley, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The used cereal flour affected the age stage-specific reproductive value (\u003cem\u003ev\u003c/em\u003e\u003csub\u003e\u003cem\u003exj\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eH. hebetor\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The earliest and highest reproductive value peaks were observed on barley (\u003cem\u003ev\u003c/em\u003e\u003csub\u003e10\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;44.60). The lowest peak of reproductive value was obtained on corn (\u003cem\u003ev\u003c/em\u003e\u003csub\u003e12\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;8.05).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003ePopulation growth parameters and Population projection.\u003c/b\u003e The third-trophic level effects of used cereal flour on the population parameters of \u003cem\u003eH.hebetor\u003c/em\u003e using a paired bootstrap test (at 5% level) are demonstrated in Table\u0026nbsp;2. The highest values of the net reproductive rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e), the intrinsic rate of increase (\u003cem\u003er\u003c/em\u003e), and the finite rate of increase (λ) were observed on corn. The effects of cereal flour on the projected population growth of \u003cem\u003eH. hebetor\u003c/em\u003e are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. The highest population size was on the corn, then followed by wheat and barley during 60 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Also, the range of variability host type in population growth indicated that the lowest degree of uncertainty in the population growth of the parasitoid was on corn (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eBiological and population traits of parasitoids depend on several factors such as, the host plant, host species and, it\u0026rsquo;s physiological condition\u003csup\u003e7,11, 18\u0026ndash;20\u003c/sup\u003e. The type of host plants involved in the developmental time period of parasitoids by shortening their preadults duration, increasing survival rate, adult body size, adult longevity, reproduction days and, fecundity\u003csup\u003e21\u003c/sup\u003e as well as increasing the population growth parameters\u003csup\u003e4\u003c/sup\u003e. This research was studied owing to the importance of host plant type on the foraging behavior, physiological and population dynamics of parasitoids\u003csup\u003e6, 22, 23\u003c/sup\u003e, and consequently to have successful production of biocontrol agents. Alongside of our previous study\u003csup\u003e16\u003c/sup\u003e, the current research was reported the third-trophic level effects of the cereal flour on the biological, life table traits, and population projection of \u003cem\u003eH. hebetor\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eOur results demonstrated that usage of corn and barley cereal flour, accelerated the immature developmental time of \u003cem\u003eH. hebetor\u003c/em\u003e, and increased the survival rate, adult longevity as well as reproduction days. Amir-Maafi \u0026amp; Chi\u003csup\u003e4\u003c/sup\u003e reported a developmental time of 10.83 d for \u003cem\u003eH. hebetor\u003c/em\u003e on the fifth instar larvae of \u003cem\u003eE. kuehniella\u003c/em\u003e fed on wheat. The preadult periods of the parasitoid in our study are within the same range as this report. However,\u003csup\u003e7\u003c/sup\u003e obtained a developmental time of 17.05 d of \u003cem\u003eH. hebetor\u003c/em\u003e on the cotton bollworm, \u003cem\u003eH. armigera\u003c/em\u003e, which fed on the leaf and fruit of tomato, it wasmore than in our study. Saadat et al. \u003csup\u003e6\u003c/sup\u003e concluded that host plants are rich in sugar and glycogen and provide more suitable physiological conditions than host species that contain terpenes and tannins for \u003cem\u003eH. hebetor.\u003c/em\u003e Our results showed that the total number of eggs/female during the lifetime of \u003cem\u003eH. hebetor\u003c/em\u003e varied from 70.87 to 91.20 eggs/female on \u003cem\u003eE. kuehniella\u003c/em\u003e, which values are much higher than the 66.30 eggs/female on \u003cem\u003eE.kuehniella\u003c/em\u003e \u003csup\u003e4\u003c/sup\u003e and 61.62 eggs/female on \u003cem\u003eH. armigera\u003c/em\u003e \u003csup\u003e7\u003c/sup\u003e. Variation in preadult developmental time and fecundity rates among these studies could be because of different host species, quality of host type, rearing conditions, and physiological condition of host species\u003csup\u003e4,6,14\u003c/sup\u003e. Seyedi et al. \u003csup\u003e14\u003c/sup\u003e stated that the highest protein content of corn (var. 704) resulted in the highest protein content and proteolytic activity of \u003cem\u003eE. kuehniella\u003c/em\u003e. Similarly, Saadat et al. \u003csup\u003e6\u003c/sup\u003e stated that the quality and quantity of protease and amylase activity of \u003cem\u003eH. hebector\u003c/em\u003e guts was greatest when reared on \u003cem\u003eE. kuehniella\u003c/em\u003e. This fact implies the importance of the quality and quantity of host type on the physiological condition at the tritrophic level among host plants, phytophagous pests, and their natural enemies\u003csup\u003e11,12\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eConstruction and analyzing the life table data based on the age-stage, two-sex life table\u003csup\u003e24,25\u003c/sup\u003e, not only consider the importance of each stages on the population dynamics,, but also it did not ignore the male population\u003csup\u003e17,26\u003c/sup\u003e. Our results demonstrated that the corn variety used in this research, was found to be more effective on \u003cem\u003eH. hebetor\u003c/em\u003e population growth rate of offspring by increasing the net reproductive rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e), the intrinsic rate of increase (\u003cem\u003er\u003c/em\u003e), the finite rate of increase (\u003cem\u003eλ\u003c/em\u003e), and shortening the mean generation time (\u003cem\u003eT\u003c/em\u003e). The highest rate of intrinsic rates of increase on corn reflected the combined effects of age-specific fecundity (\u003cem\u003em\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e) and age-specific survival rate (\u003cem\u003el\u003c/em\u003e\u003csub\u003e\u003cem\u003ex\u003c/em\u003e\u003c/sub\u003e)\u003csup\u003e27\u003c/sup\u003e. The population growth parameters of the parasitoid in our study are more than those of Amir-Maafi \u0026amp; Chi \u003csup\u003e4\u003c/sup\u003e study. These variations might be due to the variety of host types.\u003c/p\u003e \u003cp\u003eThe uncertainty based on the 2.5th and 97.5th percentiles of the net reproductive rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e) is another advantage of the age-stage, two-sex life table theory. Our results showed the same trends of population projection with population parameters. The predicted population of \u003cem\u003eH. hebetor\u003c/em\u003e was the fastest and highest on corn, followed by wheat, and barley, which was reported for the first time in this study.\u003c/p\u003e \u003cp\u003eIn conclusion, corn is a more suitable host than wheat and barley for the parasitoid, which is reared on \u003cem\u003eE. kuehniella\u003c/em\u003e. It is concluded that corn type, by providing more suitable nutrition, resulted in a shortening the developmental time, increasing the fecundity and survival rate. Interestingly, the suitability of the corn variety in this study, was confirmed once more by the estimation of population projection.\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003cp\u003e \u003cb\u003eCereal flour and insects rearing.\u003c/b\u003e The current study was designed along with our previous studies\u003csup\u003e16\u003c/sup\u003e. In this study, we used the same cereal flour including, wheat (\u003cem\u003evar\u003c/em\u003e. Homma), barley (\u003cem\u003evar\u003c/em\u003e. Makui) and, corn (var. 704), to assign how the host type affects the efficacy of parasitoid, \u003cem\u003eH. hebector\u003c/em\u003e. Aforementioned seeds were powdered using grinder and kept in a refrigerator (4 \u0026ordm;C) to avoid any contamination\u003csup\u003e28\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAccording to Tarlack et al.\u003csup\u003e28\u003c/sup\u003e method, the rearing of Mediterranean flour moth has been conducted on the aforementioned different cereal flour.\u003c/p\u003e \u003cp\u003eFor parasitoid rearing, the initial colony of \u003cem\u003eH. hebetor\u003c/em\u003e was prepared kindly from the insectarium of Agriculture Research Center, West Azerbaijan, Iran. To obtain the same aged of parasitoid\u0026rsquo;s egg, the obtained adults of \u003cem\u003eH. hebetor\u003c/em\u003e were released on the fifth instar larvae of \u003cem\u003eE.kuehniella\u003c/em\u003e, as preferred instar larvae\u003csup\u003e4\u003c/sup\u003e for 24 h. The insects rearing were continued for two generations before using in experiments at controlled condition (25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026ordm;C, 60\u0026thinsp;\u0026plusmn;\u0026thinsp;5% R. H., and a photoperiod of 14: 10 h. (L: D)).\u003c/p\u003e \u003cp\u003e \u003cb\u003eDevelopmental time and life table studies.\u003c/b\u003e To study biology and demographic parameters of \u003cem\u003eH. hebetor\u003c/em\u003e, one egg of parasitoid was left on each larva of \u003cem\u003eE. kuehniella\u003c/em\u003e and, the rest of eggs were removed\u003csup\u003e4\u003c/sup\u003e. Hence, the current study was used 49, 49 and, 58 numbers of the fifth instar parasitized larvae of \u003cem\u003eE. kuehniella\u003c/em\u003e which obtained on wheat, corn and, barley, respectively. Parasitized larvae containing one egg of \u003cem\u003eH. hebetor\u003c/em\u003e were transferred individually to plastic Petri dishes (3 cm in height, 8 cm in diameter). The developmental stage of each individual was recorded daily. By emerging adults, they were paired in plastic containers (4 cm in diameter, 10 cm in height) and fed by 10% honey solution during the oviposition period. According to our previous study\u003csup\u003e16\u003c/sup\u003e, four numbers of fifth instar larvae of \u003cem\u003eE. kuehniella\u003c/em\u003e were supplied daily to parasitize during the oviposition period. The survival, adult longevity, oviposition period and, fecundity were recorded daily. All experiments were conducted in laboratory conditions as mentioned earlier.\u003c/p\u003e \u003cp\u003e \u003cb\u003eData analysis.\u003c/b\u003e The third-trophic level effects of three different cereal flour on the developmental time, survival, and reproduction of \u003cem\u003eH. hebetor\u003c/em\u003e were analyzed according to the age-stage, two-sex life table theory \u003csup\u003e24,29\u003c/sup\u003e using the computer program TWOSEX-MSChart\u003csup\u003e30\u003c/sup\u003e. The bootstrap technique was used to estimate variances and standard errors of life history and population parameters\u003csup\u003e31,32\u003c/sup\u003e. The differences between treatments were assessed using the paired bootstrap test based on the confidence interval of differences\u003csup\u003e33,34\u003c/sup\u003e. The population growth of the parasitoid cohorts was projected according to the technique developed by Chi\u003csup\u003e35\u003c/sup\u003e using the computer program TIMING-MSChart\u003csup\u003e36\u003c/sup\u003e. The life tables based on the 0.025th and 0.975th bootstrap results of the net reproductive rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e), were used to project the uncertainty of the population growth of \u003cem\u003eH.hebetor\u003c/em\u003e\u003csup\u003e37\u003c/sup\u003e.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of Interest\u003c/h2\u003e \u003cp\u003eThere is no conflict of interest among authors.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis project was funded by Urmia University\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eN. M.and N. N. conducted the experiments. F. M.designed the experiments, analysed data, and wrote the manuscript. R. A. edited the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e \u003cp\u003eWe are appreciated to Urmia University for financial support.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eAll data generated or analysed during this study are included in this published article\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eGunduz, E. A. \u0026amp; Gulel, A. Effects of adult age and species on development period of parasitoid \u003cem\u003eBracon hebetor\u003c/em\u003e Say (Hymenoptera: Braconidae). J. Agri. Sci, 20, 31\u0026ndash;36 (2005).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTuerxun, A., Guo, W. C., Akedan, L. H. B., Xu, J. J. \u0026amp; He, J. Description of frequent species of \u003cem\u003eBraconid\u003c/em\u003e in cotton field in Xinjiang China. 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Taichung, Taiwan: National Chung Hsing University; Available from \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://140.120.197.173/Ecology/Download/TWOSEX-MSChart.rar\u003c/span\u003e\u003cspan address=\"http://140.120.197.173/Ecology/Download/TWOSEX-MSChart.rar\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2021a).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEfron, B. \u0026amp; Tibshirani, R. J. An Introduction to the Bootstrap. Chapman \u0026amp; Hall/CRC, Monographs on Statistics and Applied Probability, New York. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/978-1-4899-4541-9\u003c/span\u003e\u003cspan address=\"10.1007/978-1-4899-4541-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (1993).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang, Y. B. \u0026amp; Chi, H. 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Demography of \u003cem\u003eCacopsylla chinensis\u003c/em\u003e (Hemiptera: Psyllidae) reared on four cultivars of \u003cem\u003ePyrus bretschneideri\u003c/em\u003e and \u003cem\u003eP. communis\u003c/em\u003e (Rosales: Rosaceae) pears with estimations of confidence intervals of specific life table statistics. J. Econ Entomol, doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1093/jee/toaa149\u003c/span\u003e\u003cspan address=\"10.1093/jee/toaa149\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChi, H. Timing of control based on the stage structure of pest populations: A simulation approach. J. Econ. Entomol, 83, 1143\u0026ndash;1150 (1990).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChi, H. TIMING-MSChart: a computer program for the population projection based on age-stage, two-sex life table. Taichung, Taiwan: National Chung Hsing University; Available from http://140.120.197.173/Ecology/Download/TimingMSChart.rar (2021b).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang, H. W., Chi, H. \u0026amp; Smith, C. L. Linking demography and consumption of \u003cem\u003eHenosepilachna vigintioctopunctata\u003c/em\u003e (Coleoptera: Coccinellidae) fed on \u003cem\u003eSolanum photeinocarpum\u003c/em\u003e (Solanales: Solanaceae): With a new method to project the uncertainty of population growth and consumption. J. Econ Entomol, 111 (1), 1\u0026ndash;9 (2018).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"803\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" colspan=\"7\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTable 1.\u0026nbsp;\u003c/strong\u003eThird-trophic level effects (Mean\u0026plusmn; SE) of cereal flour on the life history parameters of \u003cem\u003eHabrobracon hebetor\u0026nbsp;\u003c/em\u003ereared on 5\u003csup\u003eth\u003c/sup\u003e instar larvae of \u003cem\u003eEphestia kuehniella\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.386277001270647%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.589580686149937%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.756035578144854%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.662007623888183%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTreatments (n\u003csup\u003e*\u003c/sup\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.015247776365946%\" colspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"5.5908513341804325%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eLife history parameters\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eWheat\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCorn\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eBarley\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\"\u003e\n \u003cp\u003eEgg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e2.00\u0026plusmn;0.00\u003csup\u003ea\u003c/sup\u003e (37)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e2.00\u0026plusmn;0.00\u003csup\u003ea\u0026nbsp;\u003c/sup\u003e(46)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e2.00\u0026plusmn;0.00\u003csup\u003ea\u003c/sup\u003e (38)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\"\u003e\n \u003cp\u003eLarvae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e1.83\u0026plusmn;0.12\u003csup\u003ea\u003c/sup\u003e (29)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e1.97\u0026plusmn;0.14\u003csup\u003ea\u003c/sup\u003e (36)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e1.88\u0026plusmn;0.13\u003csup\u003ea\u003c/sup\u003e (32)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\"\u003e\n \u003cp\u003ePupa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e7.66\u0026plusmn;0.16\u003csup\u003ea\u003c/sup\u003e (29)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e6.80\u0026plusmn;0.17\u003csup\u003eb\u003c/sup\u003e (34)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e6.84\u0026plusmn;0.16\u003csup\u003eb\u003c/sup\u003e (32)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\"\u003e\n \u003cp\u003ePre-adult\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e11.48\u0026plusmn;0.10\u003csup\u003ea\u003c/sup\u003e (29)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e10.76\u0026plusmn;0.08\u003csup\u003eb\u003c/sup\u003e (34)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e10.72\u0026plusmn;0.09\u003csup\u003ec\u003c/sup\u003e (32)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\"\u003e\n \u003cp\u003ePre-adult survival rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e0.78\u0026plusmn;0.007\u003csup\u003eb\u003c/sup\u003e (29)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e0.74\u0026plusmn;0.006\u003csup\u003ea\u003c/sup\u003e (34)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e0.84\u0026plusmn;0.006\u003csup\u003ea\u003c/sup\u003e (32)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003eFemale longevity\u003c/p\u003e\n \u003cp\u003eMale longevity\u003c/p\u003e\n \u003cp\u003eFemale total longevity\u003c/p\u003e\n \u003cp\u003eMale total longevity\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e11.00\u0026plusmn;1.03\u003csup\u003eb\u003c/sup\u003e(15)\u003c/p\u003e\n \u003cp\u003e7.57\u0026plusmn;0.81\u003csup\u003ea\u0026nbsp;\u003c/sup\u003e(14)\u003c/p\u003e\n \u003cp\u003e22.20\u0026plusmn;1.09\u003csup\u003eb\u003c/sup\u003e (15)\u003c/p\u003e\n \u003cp\u003e19.36\u0026plusmn;0.80\u003csup\u003ea\u003c/sup\u003e (14)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e14.93\u0026plusmn;0.92\u003csup\u003ea\u003c/sup\u003e(15)\u003c/p\u003e\n \u003cp\u003e5.88\u0026plusmn;0.34\u003csup\u003eb\u0026nbsp;\u003c/sup\u003e(19)\u003c/p\u003e\n \u003cp\u003e25.93\u0026plusmn;0.95\u003csup\u003ea\u0026nbsp;\u003c/sup\u003e(15)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e16.47\u0026plusmn;0.34\u003csup\u003eb\u003c/sup\u003e (19)\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e13.06\u0026plusmn;1.10 \u003csup\u003eab\u0026nbsp;\u003c/sup\u003e(15)\u003c/p\u003e\n \u003cp\u003e5.89\u0026plusmn;0.30\u003csup\u003eab\u0026nbsp;\u003c/sup\u003e(17)\u003c/p\u003e\n \u003cp\u003e23.80\u0026plusmn;1.15\u003csup\u003eab\u003c/sup\u003e (15)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e16.30\u0026plusmn;0.32\u003csup\u003eb\u003c/sup\u003e (17)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003eAPRP\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e0.33\u0026plusmn;0.13\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e0.33\u0026plusmn;0.13\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e0.20\u0026plusmn;0.11\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003eTPRP\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e11.53\u0026plusmn;0.17\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e11.33\u0026plusmn;0.20\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e10.93\u0026plusmn;0.18\u003csup\u003eb\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003eReproductive days\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e8.13\u0026plusmn;0.83\u003csup\u003eab\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e10.60\u0026plusmn;0.70\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e8.93\u0026plusmn;0.52\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.219178082191782%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"28.019925280199253%\" valign=\"top\"\u003e\n \u003cp\u003eReproduction (offspring/female)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.442092154420921%\"\u003e\n \u003cp\u003e70.87\u0026plusmn;8.23\u003csup\u003eb\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.31008717310087%\"\u003e\n \u003cp\u003e91.20\u0026plusmn;9.05\u003csup\u003ea\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.536737235367372%\" colspan=\"2\"\u003e\n \u003cp\u003e79.73\u0026plusmn;7.27\u003csup\u003eab\u003c/sup\u003e (15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.4719800747198%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eMeans in the same row followed by different letter are significantly different (paired bootstrap test at 5% significant level). Standard errors were measured by 100,000 bootstrap resampling. APRP=Adult pre-reproductive day; TPRP=Total pre-reproductive day; *n=sample size of each stage of \u003cem\u003eHabrobracon hebetor\u003c/em\u003e on each treatment\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2.\u0026nbsp;\u003c/strong\u003eThird-trophic level effects (Mean\u0026plusmn; SE) of cereal flour on the population growth parameters of \u003cem\u003eHabrobracon hebetor\u003c/em\u003e reared on 5\u003csup\u003eth\u003c/sup\u003e instar larvae of \u003cem\u003eEphestia kuehniella\u003c/em\u003e.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"682\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"30.93841642228739%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003ePopulation growth parameters \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.407624633431084%\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eWheat\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.14076246334311%\"\u003e\n \u003cp\u003e\u003cstrong\u003eTreatments (n\u003csup\u003e*\u003c/sup\u003e)\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eCorn\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.513196480938415%\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eBarley\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"30.93841642228739%\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e\u0026nbsp;R\u003csub\u003e0\u003c/sub\u003e\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e(offspring/individual) \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.407624633431084%\"\u003e\n \u003cp\u003e21.70\u0026plusmn;5.26\u003csup\u003eb\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.14076246334311%\"\u003e\n \u003cp\u003e27.92\u0026plusmn;6.57\u003csup\u003ea\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.513196480938415%\"\u003e\n \u003cp\u003e20.62\u0026plusmn;4.94\u003csup\u003eb\u003c/sup\u003e (58)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"30.93841642228739%\"\u003e\n \u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003er\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp; (day\u003csup\u003e-1\u003c/sup\u003e)\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.407624633431084%\" valign=\"top\"\u003e\n \u003cp\u003e0.2030\u0026plusmn;0.0171\u003csup\u003eb\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.14076246334311%\" valign=\"top\"\u003e\n \u003cp\u003e0.2217\u0026plusmn;0.0178\u003csup\u003ea\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.513196480938415%\" valign=\"top\"\u003e\n \u003cp\u003e0.2063\u0026plusmn;0.0178\u003csup\u003eb\u003c/sup\u003e (58)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"30.93841642228739%\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e\u0026lambda;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;(day\u003csup\u003e-1\u003c/sup\u003e)\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.407624633431084%\"\u003e\n \u003cp\u003e1.23\u0026plusmn;0.02\u003csup\u003eb\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.14076246334311%\"\u003e\n \u003cp\u003e1.25\u0026plusmn;0.02\u003csup\u003ea\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.513196480938415%\"\u003e\n \u003cp\u003e1.23\u0026plusmn;0.02\u003csup\u003eb\u003c/sup\u003e (58)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"30.93841642228739%\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eT\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;(day)\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.407624633431084%\"\u003e\n \u003cp\u003e15.16\u0026plusmn;0.38\u003csup\u003ea\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.14076246334311%\"\u003e\n \u003cp\u003e15.02\u0026plusmn;0.24\u003csup\u003e\u0026nbsp;b\u003c/sup\u003e (49)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.513196480938415%\"\u003e\n \u003cp\u003e14.66\u0026plusmn;0.22\u003csup\u003eb\u003c/sup\u003e (58)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003eThe means followed by different letters in each raw are significantly different (paired- bootstrap at 5% significance level). Standard errors were measured by 100,000 bootstrap resampling.\u003cem\u003e\u0026nbsp;R\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e = net reproductive rate; \u003cem\u003er\u003c/em\u003e = intrinsic rate of increase; \u0026lambda; = finite rate of increase; T = mean generation time; * n = sample size of each stage of \u003cem\u003eHabrobracon hebetor\u003c/em\u003e on each treatment\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"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":"Ectoparasitoid wasp, Cereal flour, Demographic parameters, Age-stage two-sex life table, Population projection","lastPublishedDoi":"10.21203/rs.3.rs-4632393/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4632393/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe parasitoid wasp, \u003cem\u003eHabracon hebetor\u003c/em\u003e, Say is used widely in mass production against larval stage of field and stored product insect pests. Less is known about the third-trophic level effects of cereal flour on the population dynamics of \u003cem\u003eH. hebetor\u003c/em\u003e. In this study, tritrophic level effects of three cereal flours (wheat, corn, and barley) on population dynamics and population projection of \u003cem\u003eH. hebetor\u003c/em\u003e were studied. Data were analyzed by constructing the age-stage, two-sex life table, to take into consideration either female or male and stage differentiation. According to our results, the used corn host type (\u003cem\u003evar\u003c/em\u003e.704) was more suitable host for \u003cem\u003eH. hebetor\u003c/em\u003e than wheat and barley due to the shortening developmental time (10.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 d), increasing survival rate (0.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006), female longevity (14.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.92 d) and fecundity (91.20\u0026thinsp;\u0026plusmn;\u0026thinsp;9.05 offspring/female). Also, the population growth parameters, including the intrinsic rate of increase (0.2217\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0178 d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), finite rate of increase (1.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 d\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), and reproductive rate (27.92\u0026thinsp;\u0026plusmn;\u0026thinsp;6.57 offspring) were the highest on corn host type. Interestingly, the predicted of population growth of \u003cem\u003eH. hebetor\u003c/em\u003e was the fastest and highest on corn, followed by wheat, and barley. The results of this study provide direction to design a comprehensive program for the mass rearing of \u003cem\u003eH. hebetor\u003c/em\u003e.\u003c/p\u003e","manuscriptTitle":"Tritrophic effects of different cereal flour on life table parameters and population projection of Habrobracon hebetor (Hymenoptera: Braconidae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-23 17:22:55","doi":"10.21203/rs.3.rs-4632393/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":"4dbdad5b-6930-472a-a6a7-7a0b5c846724","owner":[],"postedDate":"July 23rd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":34880158,"name":"Biological sciences/Biological techniques"},{"id":34880159,"name":"Biological sciences/Developmental biology"},{"id":34880160,"name":"Biological sciences/Ecology"}],"tags":[],"updatedAt":"2024-09-11T13:06:22+00:00","versionOfRecord":[],"versionCreatedAt":"2024-07-23 17:22:55","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4632393","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4632393","identity":"rs-4632393","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}