Starvation tolerance and effects on fitness of predatory mite Amblyseius orientalis

Starvation tolerance and effects on fitness of predatory mite Amblyseius orientalis | 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 Research Article Starvation tolerance and effects on fitness of predatory mite Amblyseius orientalis Fujing Sheng, Jiaxing Wei, Xianjie Wang, Endong Wang, Xuenong Xu, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5315119/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 12 Mar, 2025 Read the published version in Experimental and Applied Acarology → Version 1 posted You are reading this latest preprint version Abstract Amblyseius orientalis Ehara (Acari: Phytoseiidae) has garnered significant attention as an effective predatory mite for controlling spider mites in fruit production in China. However, despite its considerable potential for pest management, A. orientalis may face food shortages during transportation and field application. Currently, there is a lack of research on the effects of starvation on the development, survival, and reproductive capacity of A. orientalis . In this study, we evaluated the impact of starvation during both the developmental and adult stages on the development, survival, and reproduction of A. orientalis . We found that female adults exhibited greater tolerance to starvation than larvae and protonymphs. Approximately 50% of the female adults survived after eight days of starvation. Interestingly, starvation during the early developmental stages extended lifespan, nearly doubling male longevity from 20.56 to 38.00 days, and increasing female longevity from 44.68 to 70.31 days. However, starvation in female adults reduced egg production from 18.46 to 5.33 eggs over a period of ten days, while male reproductive abilities increased from 18.46 to 19.41 eggs. Additionally, the sex ratio of the offspring was not influenced by paternal starvation, but maternal starvation resulted in a male-biased offspring ratio. In conclusion, our study demonstrates that A. orientalis can tolerate starvation at various life stages, enabling it to withstand food shortages during both production and application. developmental stage hunger effect reproduction life span offspring sex ratio Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction Amblyseius orientalis Ehara (Acari: Phytoseiidae) is a widely distributed and effective predatory mite for controlling spider mites in fruit production in China (Sheng et al. 2014 ; Zheng et al. 2008 ). Initially used exclusively as a spider mite control agent (Zhang et al. 1992 ; Zheng et al. 2008 ), it has proven effective against thrips and whiteflies in the past decade as well (Sheng et al. 2014 ; Yang et al. 2018 ). Consequently, the mass rearing and large-scale production of this native predatory mite have garnered increasing interest. A. orientalis products are now employed for pest mite control in numerous orchards and greenhouses across China. However, in mass rearing, the availability and stages of alternative prey often limit A. orientalis production (Wei et al. 2023 ). Another risk factor related to prey is starvation during the storage and transportation of A. orientalis products to fields, as well as during early release for pest control. Starvation can negatively impact arthropod fitness, affecting development, longevity, and fecundity. For instance, the larvae of Hyphantria cunea and Spodoptera frugiperda pupated prematurely under starvation stress, leading to significantly reduced plumage rates and egg-laying capacity (Ju et al. 2008 ; Zeng et al. 2022 ). Conversely, varying levels of starvation can have positive effects; starved Bagrada hilaris exhibited increased responsiveness and moved farther than their non-starved counterparts (Grettenberger and Joseph, 2019 ). Additionally, appropriate starvation conditions promoted Stethorus parapauperculus to prey on Tetranychus cinnabarinus without adversely affecting physiological metabolic levels (Chen et al. 2020 ). Predatory mites often encounter prey shortages while foraging in their natural environments (Ghazy et al. 2016 ). The ability to tolerate starvation is a key indicator when selecting predators as potential commercial natural enemies (Rotheray et al. 1984). Five species of predatory mites have demonstrated a certain degree of high-temperature starvation tolerance, including Neoseiulus californicus , Amblyseius pseudolongispinosus , Amblyseius makuwa , Amblyseius eharai , and Euseius nicholsi (Qin and Li 2013 ). Among these, N. californicus exhibits particularly high tolerance to starvation. Interestingly, short-term starvation in N. californicus enhances male mating duration without significantly affecting female fertility, while prolonged starvation results in decreased male fertility (Lu et al. 2022 ). In contrast, female adults of Phytoseiulus persimilis and Iphiseius degenerans are sensitive to starvation, exhibiting reduced survival rates when deprived of food (de Courcy Williams et al. 2004 ). To understand how starvation impacts development and reproduction during production and transportation, we assessed the effects of food shortages across various life stages on A. orientalis survival and fecundity. Our study aims to provide insights into the starvation resistance of A. orientalis , which is crucial for the persistence of its populations throughout the processes of production, transportation, and release. Materials and Methods Mite rearing Amblyseius orientalis was initially collected from soybean fields in Hebei province (119°09’E, 39°43’N) and had been maintained on Carpoglyphus lactis for 10 years in the Laboratory of Predatory Mites, Institute of Plant Protection, Chinese Academy of Agricultural Sciences (Wei et al. 2023 ; Yan et al. 2024 ). All mites were reared at 25 ± 1℃, 70 ± 5% RH and L14: D10 photoperiod in the incubators (RXZ, Ningbo, China). The starvation tolerance of different stages of Amblyseius orientalis To investigate the starvation tolerance of A. orientalis at different developmental stages, larval, protonymph, and adult stages of A. orientalis were chosen for survival tests without food. The control of respective stages was reared with prey C. lactis . Every single mite was maintained in a small arena, which consisted of two layers: a transparent acrylic sheet (30 × 20 × 3 mm) with a central hole of 10 mm in diameter, sealed with nylon mesh and glue, and a rectangular glass piece (30 × 20 × 1 mm, Yan et al. 2024 ). For the starvation test on protonymphs and adults, C. lactis eggs were provided for the previous developmental stages of A. orientalis to ensure their normal development. Each stage was repeated with 40–50 A. orientalis individuals. The survival rates were recorded daily until all mites died. Effects of immature starvation on Amblyseius orientalis fitness We studied the impact of starvation during the immature stage on the development and reproduction of A. orientalis . After hatching, A. orientalis nymphs were allowed to feed on only 5 eggs of C. lactis per day, while the control group was provided with a sufficient number of C. lactis eggs. We recorded the survival rate and lifespan of both groups. Upon reaching adulthood, both starved and control individuals were given ample amounts of C. lactis . Starved A. orientalis female adults were mated with normal male adults, and starved male adults were mated with normal female adults. Normal female and male adults served as the control group. We recorded the egg production, oviposition period, and offspring sex ratio for the control group and the two treatment groups. Each treatment was repeated with 30 A. orientalis individuals. Adult fasting impact on Amblyseius orientalis reproduction A. orientalis individuals were provided with sufficient food from egg hatching to adult emergence. Both male and female adults were then individually starved for four days. We then conducted two experiments. 1) Male starvation experiment: starved A. orientalis male adults were provided with one C. lactis adult before mating with normal female mites for 24 hours. After mating, the males were removed, and the females were provided with sufficient C. lactis for further reproduction. 2) Female starvation experiment: starved A. orientalis female adults were provided with one C. lactis adult before mating with normal males for 24 hours. After mating, the males were removed, and the starved females were given one C. lactis adult every two days until death. In the control group, both normal females and males were allowed to mate for 24 hours and were subsequently provided with sufficient food for their entire lifespan. All predatory mites were individually reared and observed in separate arenas, except during the copulation period. We recorded egg production and the sex ratio of the offspring for the control group and both treatment groups, each consisting of 30 individuals. Statistical analysis Kruskal-Wallis tests was used to examine the survival rate, while Mann-Whitney U test was conducted to examine starvation effect of different stages on life parameters and fecundity of A. orientalis (α = 0.05). The general linear model (GLM) was used to analyze the developmental duration and longevity of A. orientalis based on gender and starvation. All analyses were run in SPSS v.25.0 and results were visualized in GraphPad Prism v.8.02. Results Starvation tolerance of Amblyseius orientalis Adult female mites showed greater tolerance to starvation than larvae or protonymphs ( χ 2 = 79.06, df = 2, p < 0.001; Fig. 1 ). On day 5 without prey, 30% of larvae and protonymphs had died, while all adult females survived. After seven days of starvation, all larvae and protonymphs were dead. Approximately 50% of the adult female mites succumbed by day 8. By two weeks, all adult mites had died from starvation. Effect of immature starvation on Amblyseius orientalis fitness Starvation during the immature stages affected the developmental period of both male and female A. orientalis (Fig. 2 ). The male nymph stage was extended from 2.38 to 2.89 days ( Z = -2.962, p = 0.003), while the female nymph stage increased from 2.65 to 3.90 days ( Z = -7.405, p < 0.001). Starvation also increased the total developmental period for female mites from 6.33 to 7.73 days ( Z = -5.791, p < 0.001) and for males from 20.62 to 30.42 days ( Z = 0.012, p = 0.012). Furthermore, starvation during development prolonged the total lifespan of both male and female adults (Fig. 3 ). The lifespan of male mites nearly doubled from 20.56 to 38.00 days ( Z = -5.129, p < 0.001), and female mites lifespan increased from 44.68 to 70.31 days ( Z = -5.332, p < 0.001). Statistical analyses indicated that gender, starvation, and their interaction significantly affected total lifespan (Table 1 ). Table 1 Analysis results of developmental starvation based on generalized linear model. G: gender; S: starvation; G×S: interaction of gender and starvation. Asterisks indicate statistical significance. *, p < 0.05; **, p < 0.01; ***, p < 0.001. Traits Term χ 2 value P value Developmental duration (egg) G χ 2 (1) = 80469 0.004** S χ 2 (1) < 0.0001 0.987 G×S χ 2 (1) = 0.058 0.81 Developmental duration (larva) G χ 2 (1) = 3.487 0.062 S χ 2 (1) = 2.265 0.132 G×S χ 2 (1) = 0.016 0.9 Developmental duration (nymph) G χ 2 (1) = 34.353 < 0.001*** S χ 2 (1) = 65.98 < 0.001*** G×S χ 2 (1) = 11.402 < 0.001*** Developmental duration (total) G χ 2 (1) = 9.961 0.002** S χ 2 (1) = 32.268 < 0.001*** G×S χ 2 (1) = 4.305 0.038* Longevity G χ 2 (1) = 141.687 < 0.001*** S χ 2 (1) = 82.523 < 0.001*** G×S χ 2 (1) = 2.977 0.084 Immature starvation positively affected the fecundity of A. orientalis (Fig. 4 ). It significantly enhanced the reproductive potential of both male and female adult mites. Starved male mites increased the 10-day egg production of normal female mites from 18.46 to 21.36 ( Z = -3.133, p = 0.002) and the total egg production from 30.65 to 43.45 ( Z = -2.730, p = 0.006). Similarly, starved female mites increased their 10-day egg production from 18.46 to 20.44 ( Z = -2.186, p = 0.029), with total egg production rising from 30.85 to 41.5 ( Z = -2.574, p = 0.010). Effect of adult starvation on Amblyseius orientalis fecundity Starvation during the adult stage significantly affected the egg-laying of A. orientalis (Figs. 7 and 8 ). Egg production in starved female mites dropped significantly from 18.46 to 5.33 eggs over 10 days ( Z = -4.995, p < 0.001; Fig. 7 ), whereas starved male mites showed an increase from 18.46 to 19.41 eggs ( Z = -3.142, p = 0.002). Additionally, the sex ratio of offspring from starved maternal parents decreased significantly from 2.01 to 0.96 ( Z = -4.060, p < 0.001; Fig. 8 ). In contrast, paternal starvation did not affect the offspring sex ratio. Discussion In this study, we discovered that adult female A. orientalis demonstrated the highest tolerance to starvation compared to other developmental stages, such as larvae and protonymphs. Remarkably, over 50% of these female adults were able to survive for a week without food. This resilience suggested that adult females possess a significant advantage in environments with limited food availability. This ability to withstand starvation enabled female adults a critical stage during the transportation and application processes. Their resilience also offered a substantial buffer period, allowing them to survive from the time they leave the production facility until they reach their destination in the field. This enhanced survival rate ultimately contributes to the successful establishment and proliferation of predatory mites, which are essential for controlling pest outbreaks. Their ability to thrive despite temporary food shortages ensures they can fulfill their role in pest management, providing a natural and sustainable solution to agricultural challenges. This characteristic underscores the importance of considering female adult mites in strategies aimed at maximizing the efficacy of biological pest control. In some cases, starvation during development positively affects the reproduction and survival capabilities. In this study, developmental starvation increased egg production by extending the egg-laying period of A. orientalis . Similarly, N. californicus experienced a prolonged pre-oviposition period after starvation (Lu et al. 2022 ). This positive effect may result from a strategy of slowing growth to conserve energy, helping them survive in resource-scarce environments and leading to a longer lifespan. For example, Cnaphalocrocis medinalis larvae extended their larval stage and increased feeding time by reducing their growth rate under starvation (Yang et al. 2015 ). However, not all predatory mites follow this strategy. P. persimilis and N. californicus accelerated their development when prey is scarce. This difference may relate to physiological mechanisms and ecological adaptations (Walzer and Schausberger 2011 ). This effect persisted into the adult stage with adequate food. A stable prey supply during P. persimilis 's developmental stage promotes optimal body size (Han et al. 2022 ). In our study, early starvation prolonged the oviposition period of adult mites with the increased egg production. Interestingly, nutrient-limited female mosquitoes laid fewer eggs, but mating with well-nourished males increased lipid investment in eggs (Zirbel et al. 2021). Additionally, developmental starvation did not significantly affect sex ratios, suggesting limited impact of early development and nutrition on sex determination (Takano and Takasu 2019 ). This adaptive strategy allowed A. orientalis to thrive in resource-limited environments, providing a survival advantage. Reducing metabolic costs under starvation stress decreased endogenous fuel use and increased survival capability (Secor and Carey 2016 ). Starvation during development may enhance resource absorption and metabolism in A. orientalis . After starvation, when adults received sufficient resources, they efficiently utilized these nutrients to maintain and promote physiological indices through aggressive feeding. This was supported by Cacosceles newmannii larvae, which adapted to food shortages and rapidly processed food when available (Smit et al. 2021 ). This resource usage enabled starved A. orientalis to nearly double their lifespan. Similarly, food scarcity increased lifespan in Tetranychus urticae and Neoseiulus cucumeris (Lee et al. 2020 ; Li and Zhang 2022 ), and boosted fecundity in Drosophila melanogaster (Zajitschek et al. 2016 ). These findings emphasized that starvation, as a biotic stressor, profoundly affected physiological and ecological adaptations in A. orientalis . However, starvation during the adult stage negatively impacted A. orientalis reproduction. Specifically, 10-day egg production of female adults significantly decreased, indicating they cannot sustain reproductive requirements under food shortage. P. persimilis also showed reduced parental nutrition lowered offspring survival (Han et al. 2024 ). Additionally, a significant decreased in the female-to-male offspring ratio among starved females indicated nutritional shortages for gravid mothers. Starvation was reported to impact offspring sex ratios in predatory mite species. As prey T. urticae decreased, female offspring proportions of P. persimilis and Amblyseius womersley dropped significantly (Toyoshima and Amano 1998 ). Under resource limitations, the higher energy demands of female development challenged fertilized eggs, resulting in more male offspring. This shift in sex ratio indicated food scarcity by providing one adult of C. lactis . Our study confirmed the delicate balance between reproduction and starvation in adults, allowing predatory mite populations to survive with limited food supply. A. orientalis showed a preference for the developmental stage of alternative prey C. lactis during rearing (Wei et al. 2023 ). To support energy needs for growth, development, and reproduction, it is vital to have sufficient prey, particularly young stages, to sustain stable A. orientalis reproduction. In mass rearing, maintaining a balanced mix of prey at various stages is thus crucial for efficient production. Strategies such as reducing C. lactis adults to provide space for younger prey help keep immature populations dominant. These measures are essential for successful mass rearing, as unfavorable prey stages result in inadequate energy for growth. In summary, the starvation tolerance of A. orientalis during various life stages buffered food shortage challenges during production and transportation as commercialized natural enemies. This capability extends their shelf life and storage duration, enhancing effectiveness in biological control programs. A large prey supply was found unnecessary for mass rearing, as it increased costs and reduces efficiency. Understanding starvation tolerance promotes stable growth and reproduction in A. orientalis production, boosting their reproductive potential in the field for pest control. Declarations Competing Interests The authors have declared that no competing interest exist. Author Contribution F.S.: Investigation and Formal analysis. J.W.: Formal analysis, Original draft preparation, Review and Editing. E.W.: Validation, Supervision and Editing. X.W.: Validation, Supervision and Editing. X.X.: Validation, Supervision and Editing. B.Z.: Conceptualization, Supervision, Funding, Original draft preparation, Review and Editing. Acknowledgments This work was supported by National Key R&D Program of China (2023YFD1400600). References Chen JY, Wang JY, Zhang FP, Li L, Han DY, Zhang CH, Fu YG (2020) Predation of Stethorus (Allosstethorus) parapauperculus in different extent of starvation to Tetranychus cinnabarinus . J Environ Entomol 42(05):1201–1209. https://doi.org/10.3969/i.issn.1674-0858.2020.05.20 de Courcy Williams ME, Kravar-Garde L, Fenlon JS, Sunderland KD (2004) The relationship between dietary specialism and availability of food and water on cannibalistic interactions among predatory mites in protected crops. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5315119","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":373809657,"identity":"7996d773-1886-49e5-a733-cd001acb9481","order_by":0,"name":"Fujing Sheng","email":"","orcid":"","institution":"Shandongzhongke Beneficial Insect Resources Utilization Technology Innovation Center","correspondingAuthor":false,"prefix":"","firstName":"Fujing","middleName":"","lastName":"Sheng","suffix":""},{"id":373809659,"identity":"5fd22736-d2df-403a-96a3-4e98a5e9db7e","order_by":1,"name":"Jiaxing Wei","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Jiaxing","middleName":"","lastName":"Wei","suffix":""},{"id":373809661,"identity":"9a9de81b-1887-41cc-92c8-33904672165d","order_by":2,"name":"Xianjie Wang","email":"","orcid":"","institution":"Shandongzhongke Beneficial Insect Resources Utilization Technology Innovation Center","correspondingAuthor":false,"prefix":"","firstName":"Xianjie","middleName":"","lastName":"Wang","suffix":""},{"id":373809663,"identity":"3fd34b01-8dc4-4ffb-a2f4-f2185839106b","order_by":3,"name":"Endong Wang","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Endong","middleName":"","lastName":"Wang","suffix":""},{"id":373809666,"identity":"66c961df-3441-4827-b07a-b5facd890aa0","order_by":4,"name":"Xuenong Xu","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Xuenong","middleName":"","lastName":"Xu","suffix":""},{"id":373809668,"identity":"f6cec7f4-5195-47c8-9abb-90c75fe820c6","order_by":5,"name":"Bo Zhang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA0ElEQVRIiWNgGAWjYDACCSBmbGDgsW9vIFGLnAHPARK1GBtIJBCpQ35287OHX3fYJG6XfPzww48/DPLmhLQwzjlmbix7Ji1x5+w0Y8neNgbDnQ0EtDBLJJhJS7YdTmy4ncMgwdvAkGBwgIAWNon0bxAtN88w//zzhwgtPBI5ZpIf2w4bG9zgYZPmYSNCi4RETpk045k0OcmeNDNr2TYJww2EtMjPSN8m+XOHDQ8/++HHN9/8sZEnaAsIMPMg2UqEeiBg/EGculEwCkbBKBipAACSmD9DQxB8VQAAAABJRU5ErkJggg==","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":true,"prefix":"","firstName":"Bo","middleName":"","lastName":"Zhang","suffix":""}],"badges":[],"createdAt":"2024-10-23 02:53:17","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5315119/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5315119/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10493-025-01008-8","type":"published","date":"2025-03-12T15:58:27+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":68356663,"identity":"18186152-a488-4f91-9869-7eb3c57f3700","added_by":"auto","created_at":"2024-11-06 11:27:47","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":124075,"visible":true,"origin":"","legend":"\u003cp\u003eSurvival curves of three developmental stages of \u003cem\u003eA. orientalis\u003c/em\u003e in the absence of prey. Asterisks indicate statistical significance. ***, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/f65c608cf35b164c1419427a.png"},{"id":68356662,"identity":"faea78be-8529-414d-a2c8-e8555ce9adba","added_by":"auto","created_at":"2024-11-06 11:27:47","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":128761,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of developmental periods of the starved males and females of \u003cem\u003eA. orientalis\u003c/em\u003e. Error bars indicate mean (±SEM). “ns” indicates non-significant. Asterisks indicate statistical significance. *, \u003cem\u003ep\u003c/em\u003e\u0026lt; 0.05; **, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01; ***, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/aad3e0b7a0cbf19d08cc4764.png"},{"id":68357011,"identity":"029a0d75-c722-4636-9bbc-ed88d8147502","added_by":"auto","created_at":"2024-11-06 11:35:47","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":34753,"visible":true,"origin":"","legend":"\u003cp\u003eThe longevity of\u003cem\u003e A. orientalis\u003c/em\u003e female and male after immature starvation. Error bars indicate mean (±SEM). Asterisks indicate statistical significance. ***, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/824bcbb8f6f5540bb5389482.png"},{"id":68357010,"identity":"276c1053-50c4-49f1-a69b-4dc1d7f32b66","added_by":"auto","created_at":"2024-11-06 11:35:47","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":111305,"visible":true,"origin":"","legend":"\u003cp\u003eEgg production of either male or female adults experiencing immature starvation. Error bars indicate mean (±SEM). “ns” indicates non-significant. Asterisks indicate statistical significance. *, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05; **, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01; ***, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/83bdad70560719e09e7669c2.png"},{"id":68356666,"identity":"d2207c79-781e-44eb-bdd3-b66c620962ff","added_by":"auto","created_at":"2024-11-06 11:27:47","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":134472,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of oviposition period time after immature starvation. Error bars indicate mean (±SEM). “ns” indicates non-significant. Asterisks indicate statistical significance. *, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/50df6edf6c1481764b54dd2e.png"},{"id":68356667,"identity":"97a16632-da35-4446-b935-012d1acc0a7e","added_by":"auto","created_at":"2024-11-06 11:27:47","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":85192,"visible":true,"origin":"","legend":"\u003cp\u003eSex ratios of the offspring produced by either starved female and male parents. Error bars indicate mean (±SEM). “ns” indicates non-significant.\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/7c0d656c2dd28c239d98130f.png"},{"id":68356670,"identity":"23bba711-dfa1-4e1d-b73f-d9ce09006088","added_by":"auto","created_at":"2024-11-06 11:27:48","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":88583,"visible":true,"origin":"","legend":"\u003cp\u003eThe number of offspring from starved parents in the first 10 days. Error bars indicate mean (±SEM). “ns” indicates non-significant. Asterisks indicate statistical significance. **, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01.\u003c/p\u003e","description":"","filename":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/849bd7705e5c1f75273a9a1d.png"},{"id":68356665,"identity":"e20c2f1a-615f-47b4-b4c8-160bd3f4ba13","added_by":"auto","created_at":"2024-11-06 11:27:47","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":96940,"visible":true,"origin":"","legend":"\u003cp\u003eSex ratios of the offspring produced by either starved males or female parents at the adult stage. Error bars indicate mean (±SEM). “ns” indicates non-significant. Asterisks indicate statistical significance. ***, \u003cem\u003ep\u003c/em\u003e\u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage8.png","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/d739015e272cfde58780266d.png"},{"id":78689081,"identity":"d507c52c-dd43-4517-9cbd-8774631efa89","added_by":"auto","created_at":"2025-03-17 16:10:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1463944,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5315119/v1/d003361e-ff99-4829-822b-b512a4d66376.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Starvation tolerance and effects on fitness of predatory mite Amblyseius orientalis","fulltext":[{"header":"Introduction","content":"\u003cp\u003e \u003cem\u003eAmblyseius orientalis\u003c/em\u003e Ehara (Acari: Phytoseiidae) is a widely distributed and effective predatory mite for controlling spider mites in fruit production in China (Sheng et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zheng et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Initially used exclusively as a spider mite control agent (Zhang et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Zheng et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), it has proven effective against thrips and whiteflies in the past decade as well (Sheng et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Consequently, the mass rearing and large-scale production of this native predatory mite have garnered increasing interest. \u003cem\u003eA. orientalis\u003c/em\u003e products are now employed for pest mite control in numerous orchards and greenhouses across China. However, in mass rearing, the availability and stages of alternative prey often limit \u003cem\u003eA. orientalis\u003c/em\u003e production (Wei et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAnother risk factor related to prey is starvation during the storage and transportation of \u003cem\u003eA. orientalis\u003c/em\u003e products to fields, as well as during early release for pest control. Starvation can negatively impact arthropod fitness, affecting development, longevity, and fecundity. For instance, the larvae of \u003cem\u003eHyphantria cunea\u003c/em\u003e and \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e pupated prematurely under starvation stress, leading to significantly reduced plumage rates and egg-laying capacity (Ju et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Zeng et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Conversely, varying levels of starvation can have positive effects; starved \u003cem\u003eBagrada hilaris\u003c/em\u003e exhibited increased responsiveness and moved farther than their non-starved counterparts (Grettenberger and Joseph, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Additionally, appropriate starvation conditions promoted \u003cem\u003eStethorus parapauperculus\u003c/em\u003e to prey on \u003cem\u003eTetranychus cinnabarinus\u003c/em\u003e without adversely affecting physiological metabolic levels (Chen et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Predatory mites often encounter prey shortages while foraging in their natural environments (Ghazy et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe ability to tolerate starvation is a key indicator when selecting predators as potential commercial natural enemies (Rotheray et al. 1984). Five species of predatory mites have demonstrated a certain degree of high-temperature starvation tolerance, including \u003cem\u003eNeoseiulus californicus\u003c/em\u003e, \u003cem\u003eAmblyseius pseudolongispinosus\u003c/em\u003e, \u003cem\u003eAmblyseius makuwa\u003c/em\u003e, \u003cem\u003eAmblyseius eharai\u003c/em\u003e, and \u003cem\u003eEuseius nicholsi\u003c/em\u003e (Qin and Li \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Among these, \u003cem\u003eN. californicus\u003c/em\u003e exhibits particularly high tolerance to starvation. Interestingly, short-term starvation in \u003cem\u003eN. californicus\u003c/em\u003e enhances male mating duration without significantly affecting female fertility, while prolonged starvation results in decreased male fertility (Lu et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In contrast, female adults of \u003cem\u003ePhytoseiulus persimilis\u003c/em\u003e and \u003cem\u003eIphiseius degenerans\u003c/em\u003e are sensitive to starvation, exhibiting reduced survival rates when deprived of food (de Courcy Williams et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo understand how starvation impacts development and reproduction during production and transportation, we assessed the effects of food shortages across various life stages on \u003cem\u003eA. orientalis\u003c/em\u003e survival and fecundity. Our study aims to provide insights into the starvation resistance of \u003cem\u003eA. orientalis\u003c/em\u003e, which is crucial for the persistence of its populations throughout the processes of production, transportation, and release.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eMite rearing\u003c/h2\u003e \u003cp\u003e \u003cem\u003eAmblyseius orientalis\u003c/em\u003e was initially collected from soybean fields in Hebei province (119\u0026deg;09\u0026rsquo;E, 39\u0026deg;43\u0026rsquo;N) and had been maintained on \u003cem\u003eCarpoglyphus lactis\u003c/em\u003e for 10 years in the Laboratory of Predatory Mites, Institute of Plant Protection, Chinese Academy of Agricultural Sciences (Wei et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Yan et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). All mites were reared at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1℃, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and L14: D10 photoperiod in the incubators (RXZ, Ningbo, China).\u003c/p\u003e \u003cp\u003e \u003cb\u003eThe starvation tolerance of different stages of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo investigate the starvation tolerance of \u003cem\u003eA. orientalis\u003c/em\u003e at different developmental stages, larval, protonymph, and adult stages of \u003cem\u003eA. orientalis\u003c/em\u003e were chosen for survival tests without food. The control of respective stages was reared with prey \u003cem\u003eC. lactis\u003c/em\u003e. Every single mite was maintained in a small arena, which consisted of two layers: a transparent acrylic sheet (30 \u0026times; 20 \u0026times; 3 mm) with a central hole of 10 mm in diameter, sealed with nylon mesh and glue, and a rectangular glass piece (30 \u0026times; 20 \u0026times; 1 mm, Yan et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). For the starvation test on protonymphs and adults, \u003cem\u003eC. lactis\u003c/em\u003e eggs were provided for the previous developmental stages of \u003cem\u003eA. orientalis\u003c/em\u003e to ensure their normal development. Each stage was repeated with 40\u0026ndash;50 \u003cem\u003eA. orientalis\u003c/em\u003e individuals. The survival rates were recorded daily until all mites died.\u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of immature starvation on\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e \u003cb\u003efitness\u003c/b\u003e\u003c/p\u003e \u003cp\u003eWe studied the impact of starvation during the immature stage on the development and reproduction of \u003cem\u003eA. orientalis\u003c/em\u003e. After hatching, \u003cem\u003eA. orientalis\u003c/em\u003e nymphs were allowed to feed on only 5 eggs of \u003cem\u003eC. lactis\u003c/em\u003e per day, while the control group was provided with a sufficient number of \u003cem\u003eC. lactis\u003c/em\u003e eggs. We recorded the survival rate and lifespan of both groups. Upon reaching adulthood, both starved and control individuals were given ample amounts of \u003cem\u003eC. lactis\u003c/em\u003e. Starved \u003cem\u003eA. orientalis\u003c/em\u003e female adults were mated with normal male adults, and starved male adults were mated with normal female adults. Normal female and male adults served as the control group. We recorded the egg production, oviposition period, and offspring sex ratio for the control group and the two treatment groups. Each treatment was repeated with 30 \u003cem\u003eA. orientalis\u003c/em\u003e individuals.\u003c/p\u003e \u003cp\u003e \u003cb\u003eAdult fasting impact on\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e \u003cb\u003ereproduction\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003cem\u003eA. orientalis\u003c/em\u003e individuals were provided with sufficient food from egg hatching to adult emergence. Both male and female adults were then individually starved for four days. We then conducted two experiments. 1) Male starvation experiment: starved \u003cem\u003eA. orientalis\u003c/em\u003e male adults were provided with one \u003cem\u003eC. lactis\u003c/em\u003e adult before mating with normal female mites for 24 hours. After mating, the males were removed, and the females were provided with sufficient \u003cem\u003eC. lactis\u003c/em\u003e for further reproduction. 2) Female starvation experiment: starved \u003cem\u003eA. orientalis\u003c/em\u003e female adults were provided with one \u003cem\u003eC. lactis\u003c/em\u003e adult before mating with normal males for 24 hours. After mating, the males were removed, and the starved females were given one \u003cem\u003eC. lactis\u003c/em\u003e adult every two days until death. In the control group, both normal females and males were allowed to mate for 24 hours and were subsequently provided with sufficient food for their entire lifespan. All predatory mites were individually reared and observed in separate arenas, except during the copulation period. We recorded egg production and the sex ratio of the offspring for the control group and both treatment groups, each consisting of 30 individuals.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eKruskal-Wallis tests was used to examine the survival rate, while Mann-Whitney U test was conducted to examine starvation effect of different stages on life parameters and fecundity of \u003cem\u003eA. orientalis\u003c/em\u003e (α\u0026thinsp;=\u0026thinsp;0.05). The general linear model (GLM) was used to analyze the developmental duration and longevity of \u003cem\u003eA. orientalis\u003c/em\u003e based on gender and starvation. All analyses were run in SPSS v.25.0 and results were visualized in GraphPad Prism v.8.02.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003e \u003cb\u003eStarvation tolerance of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eAdult female mites showed greater tolerance to starvation than larvae or protonymphs (\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;79.06, df\u0026thinsp;=\u0026thinsp;2, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). On day 5 without prey, 30% of larvae and protonymphs had died, while all adult females survived. After seven days of starvation, all larvae and protonymphs were dead. Approximately 50% of the adult female mites succumbed by day 8. By two weeks, all adult mites had died from starvation.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eEffect of immature starvation on\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e \u003cb\u003efitness\u003c/b\u003e\u003c/p\u003e \u003cp\u003eStarvation during the immature stages affected the developmental period of both male and female \u003cem\u003eA. orientalis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The male nymph stage was extended from 2.38 to 2.89 days (\u003cem\u003eZ\u003c/em\u003e = -2.962, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.003), while the female nymph stage increased from 2.65 to 3.90 days (\u003cem\u003eZ\u003c/em\u003e = -7.405, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Starvation also increased the total developmental period for female mites from 6.33 to 7.73 days (\u003cem\u003eZ\u003c/em\u003e = -5.791, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and for males from 20.62 to 30.42 days (\u003cem\u003eZ\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.012, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.012). Furthermore, starvation during development prolonged the total lifespan of both male and female adults (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The lifespan of male mites nearly doubled from 20.56 to 38.00 days (\u003cem\u003eZ\u003c/em\u003e = -5.129, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and female mites lifespan increased from 44.68 to 70.31 days (\u003cem\u003eZ\u003c/em\u003e = -5.332, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Statistical analyses indicated that gender, starvation, and their interaction significantly affected total lifespan (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAnalysis results of developmental starvation based on generalized linear model. G: gender; S: starvation; G\u0026times;S: interaction of gender and starvation. Asterisks indicate statistical significance. *, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05; **, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01; ***, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTraits\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTerm\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eP value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eDevelopmental duration (egg)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;80469\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.004**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.987\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.058\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.81\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eDevelopmental duration (larva)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;3.487\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.062\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;2.265\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.132\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.016\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eDevelopmental duration (nymph)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;34.353\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;65.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;11.402\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eDevelopmental duration (total)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;9.961\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.002**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;32.268\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;4.305\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.038*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eLongevity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;141.687\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;82.523\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eG\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eχ\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e\u003csub\u003e(1)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;2.977\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.084\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eImmature starvation positively affected the fecundity of \u003cem\u003eA. orientalis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). It significantly enhanced the reproductive potential of both male and female adult mites. Starved male mites increased the 10-day egg production of normal female mites from 18.46 to 21.36 (\u003cem\u003eZ\u003c/em\u003e = -3.133, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.002) and the total egg production from 30.65 to 43.45 (\u003cem\u003eZ\u003c/em\u003e = -2.730, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.006). Similarly, starved female mites increased their 10-day egg production from 18.46 to 20.44 (\u003cem\u003eZ\u003c/em\u003e = -2.186, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.029), with total egg production rising from 30.85 to 41.5 (\u003cem\u003eZ\u003c/em\u003e = -2.574, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.010).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eEffect of adult starvation on\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e \u003cb\u003efecundity\u003c/b\u003e\u003c/p\u003e \u003cp\u003eStarvation during the adult stage significantly affected the egg-laying of \u003cem\u003eA. orientalis\u003c/em\u003e (Figs.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e and \u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). Egg production in starved female mites dropped significantly from 18.46 to 5.33 eggs over 10 days (\u003cem\u003eZ\u003c/em\u003e = -4.995, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e), whereas starved male mites showed an increase from 18.46 to 19.41 eggs (\u003cem\u003eZ\u003c/em\u003e = -3.142, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.002). Additionally, the sex ratio of offspring from starved maternal parents decreased significantly from 2.01 to 0.96 (\u003cem\u003eZ\u003c/em\u003e = -4.060, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). In contrast, paternal starvation did not affect the offspring sex ratio.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we discovered that adult female \u003cem\u003eA. orientalis\u003c/em\u003e demonstrated the highest tolerance to starvation compared to other developmental stages, such as larvae and protonymphs. Remarkably, over 50% of these female adults were able to survive for a week without food. This resilience suggested that adult females possess a significant advantage in environments with limited food availability. This ability to withstand starvation enabled female adults a critical stage during the transportation and application processes. Their resilience also offered a substantial buffer period, allowing them to survive from the time they leave the production facility until they reach their destination in the field. This enhanced survival rate ultimately contributes to the successful establishment and proliferation of predatory mites, which are essential for controlling pest outbreaks. Their ability to thrive despite temporary food shortages ensures they can fulfill their role in pest management, providing a natural and sustainable solution to agricultural challenges. This characteristic underscores the importance of considering female adult mites in strategies aimed at maximizing the efficacy of biological pest control.\u003c/p\u003e \u003cp\u003eIn some cases, starvation during development positively affects the reproduction and survival capabilities. In this study, developmental starvation increased egg production by extending the egg-laying period of \u003cem\u003eA. orientalis\u003c/em\u003e. Similarly, \u003cem\u003eN. californicus\u003c/em\u003e experienced a prolonged pre-oviposition period after starvation (Lu et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This positive effect may result from a strategy of slowing growth to conserve energy, helping them survive in resource-scarce environments and leading to a longer lifespan. For example, \u003cem\u003eCnaphalocrocis medinalis\u003c/em\u003e larvae extended their larval stage and increased feeding time by reducing their growth rate under starvation (Yang et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). However, not all predatory mites follow this strategy. \u003cem\u003eP. persimilis\u003c/em\u003e and \u003cem\u003eN. californicus\u003c/em\u003e accelerated their development when prey is scarce. This difference may relate to physiological mechanisms and ecological adaptations (Walzer and Schausberger \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). This effect persisted into the adult stage with adequate food. A stable prey supply during \u003cem\u003eP. persimilis\u003c/em\u003e's developmental stage promotes optimal body size (Han et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In our study, early starvation prolonged the oviposition period of adult mites with the increased egg production. Interestingly, nutrient-limited female mosquitoes laid fewer eggs, but mating with well-nourished males increased lipid investment in eggs (Zirbel et al. 2021). Additionally, developmental starvation did not significantly affect sex ratios, suggesting limited impact of early development and nutrition on sex determination (Takano and Takasu \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis adaptive strategy allowed \u003cem\u003eA. orientalis\u003c/em\u003e to thrive in resource-limited environments, providing a survival advantage. Reducing metabolic costs under starvation stress decreased endogenous fuel use and increased survival capability (Secor and Carey \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Starvation during development may enhance resource absorption and metabolism in \u003cem\u003eA. orientalis\u003c/em\u003e. After starvation, when adults received sufficient resources, they efficiently utilized these nutrients to maintain and promote physiological indices through aggressive feeding. This was supported by \u003cem\u003eCacosceles newmannii\u003c/em\u003e larvae, which adapted to food shortages and rapidly processed food when available (Smit et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This resource usage enabled starved \u003cem\u003eA. orientalis\u003c/em\u003e to nearly double their lifespan. Similarly, food scarcity increased lifespan in \u003cem\u003eTetranychus urticae\u003c/em\u003e and \u003cem\u003eNeoseiulus cucumeris\u003c/em\u003e (Lee et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Li and Zhang \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and boosted fecundity in \u003cem\u003eDrosophila melanogaster\u003c/em\u003e (Zajitschek et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). These findings emphasized that starvation, as a biotic stressor, profoundly affected physiological and ecological adaptations in \u003cem\u003eA. orientalis\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eHowever, starvation during the adult stage negatively impacted \u003cem\u003eA. orientalis\u003c/em\u003e reproduction. Specifically, 10-day egg production of female adults significantly decreased, indicating they cannot sustain reproductive requirements under food shortage. \u003cem\u003eP. persimilis\u003c/em\u003e also showed reduced parental nutrition lowered offspring survival (Han et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Additionally, a significant decreased in the female-to-male offspring ratio among starved females indicated nutritional shortages for gravid mothers. Starvation was reported to impact offspring sex ratios in predatory mite species. As prey \u003cem\u003eT. urticae\u003c/em\u003e decreased, female offspring proportions of \u003cem\u003eP. persimilis\u003c/em\u003e and \u003cem\u003eAmblyseius womersley\u003c/em\u003e dropped significantly (Toyoshima and Amano \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Under resource limitations, the higher energy demands of female development challenged fertilized eggs, resulting in more male offspring. This shift in sex ratio indicated food scarcity by providing one adult of \u003cem\u003eC. lactis\u003c/em\u003e. Our study confirmed the delicate balance between reproduction and starvation in adults, allowing predatory mite populations to survive with limited food supply.\u003c/p\u003e \u003cp\u003e \u003cem\u003eA. orientalis\u003c/em\u003e showed a preference for the developmental stage of alternative prey \u003cem\u003eC. lactis\u003c/em\u003e during rearing (Wei et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). To support energy needs for growth, development, and reproduction, it is vital to have sufficient prey, particularly young stages, to sustain stable \u003cem\u003eA. orientalis\u003c/em\u003e reproduction. In mass rearing, maintaining a balanced mix of prey at various stages is thus crucial for efficient production. Strategies such as reducing \u003cem\u003eC. lactis\u003c/em\u003e adults to provide space for younger prey help keep immature populations dominant. These measures are essential for successful mass rearing, as unfavorable prey stages result in inadequate energy for growth.\u003c/p\u003e \u003cp\u003eIn summary, the starvation tolerance of \u003cem\u003eA. orientalis\u003c/em\u003e during various life stages buffered food shortage challenges during production and transportation as commercialized natural enemies. This capability extends their shelf life and storage duration, enhancing effectiveness in biological control programs. A large prey supply was found unnecessary for mass rearing, as it increased costs and reduces efficiency. Understanding starvation tolerance promotes stable growth and reproduction in \u003cem\u003eA. orientalis\u003c/em\u003e production, boosting their reproductive potential in the field for pest control.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting Interests\u003c/h2\u003e \u003cp\u003eThe authors have declared that no competing interest exist.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eF.S.: Investigation and Formal analysis. J.W.: Formal analysis, Original draft preparation, Review and Editing. E.W.: Validation, Supervision and Editing. X.W.: Validation, Supervision and Editing. X.X.: Validation, Supervision and Editing. B.Z.: Conceptualization, Supervision, Funding, Original draft preparation, Review and Editing.\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eThis work was supported by National Key R\u0026amp;D Program of China (2023YFD1400600).\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eChen JY, Wang JY, Zhang FP, Li L, Han DY, Zhang CH, Fu YG (2020) Predation of \u003cem\u003eStethorus\u003c/em\u003e (Allosstethorus) \u003cem\u003eparapauperculus\u003c/em\u003e in different extent of starvation to \u003cem\u003eTetranychus cinnabarinus\u003c/em\u003e. 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Acta Ecol Sin 28:4830\u0026ndash;4840\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZirbel K, Alto BW (2021) Paternal and maternal effects in a mosquito: A bridge for life history transition. J Insect Physiol 131:104243. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jinsphys.2021.104243\u003c/span\u003e\u003cspan address=\"10.1016/j.jinsphys.2021.104243\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":true,"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":"developmental stage, hunger effect, reproduction, life span, offspring sex ratio","lastPublishedDoi":"10.21203/rs.3.rs-5315119/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5315119/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eAmblyseius orientalis\u003c/em\u003e Ehara (Acari: Phytoseiidae) has garnered significant attention as an effective predatory mite for controlling spider mites in fruit production in China. However, despite its considerable potential for pest management, \u003cem\u003eA. orientalis\u003c/em\u003e may face food shortages during transportation and field application. Currently, there is a lack of research on the effects of starvation on the development, survival, and reproductive capacity of \u003cem\u003eA. orientalis\u003c/em\u003e. In this study, we evaluated the impact of starvation during both the developmental and adult stages on the development, survival, and reproduction of \u003cem\u003eA. orientalis\u003c/em\u003e. We found that female adults exhibited greater tolerance to starvation than larvae and protonymphs. Approximately 50% of the female adults survived after eight days of starvation. Interestingly, starvation during the early developmental stages extended lifespan, nearly doubling male longevity from 20.56 to 38.00 days, and increasing female longevity from 44.68 to 70.31 days. However, starvation in female adults reduced egg production from 18.46 to 5.33 eggs over a period of ten days, while male reproductive abilities increased from 18.46 to 19.41 eggs. Additionally, the sex ratio of the offspring was not influenced by paternal starvation, but maternal starvation resulted in a male-biased offspring ratio. In conclusion, our study demonstrates that \u003cem\u003eA. orientalis\u003c/em\u003e can tolerate starvation at various life stages, enabling it to withstand food shortages during both production and application.\u003c/p\u003e","manuscriptTitle":"Starvation tolerance and effects on fitness of predatory mite Amblyseius orientalis","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-11-06 11:27:43","doi":"10.21203/rs.3.rs-5315119/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":"551d871e-f259-42c7-a4c4-d933c63896b3","owner":[],"postedDate":"November 6th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-03-17T16:04:24+00:00","versionOfRecord":{"articleIdentity":"rs-5315119","link":"https://doi.org/10.1007/s10493-025-01008-8","journal":{"identity":"experimental-and-applied-acarology","isVorOnly":false,"title":"Experimental and Applied Acarology"},"publishedOn":"2025-03-12 15:58:27","publishedOnDateReadable":"March 12th, 2025"},"versionCreatedAt":"2024-11-06 11:27:43","video":"","vorDoi":"10.1007/s10493-025-01008-8","vorDoiUrl":"https://doi.org/10.1007/s10493-025-01008-8","workflowStages":[]},"version":"v1","identity":"rs-5315119","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5315119","identity":"rs-5315119","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}