Both heat stress and prey species affect Amblyseius orientalis performance

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Both temperature and prey closely impact the fitness and predation of natural enemies during biological control. Amblyseius orientalis (Ehara) (Acari: Phytoseiidae) is a native and effective predatory mite against spider mites and small sucking insects in many kinds of orchards and greenhouses, where high temperatures always occur during hot summer. However, the effects of heat stress and prey species on the fitness and predatory efficiency of A. orientalis have not been well understood. In current study, we selected two preys of Tetranychus urticae and Bemisia tabaci and two temperatures of 25℃ and 33℃ to explore the performances of A. orientalis under four combination treatments. The results showed that both temperature and prey significantly affect A. orientalis fitness. In detail, A. orientalis had the longest developmental duration of 7.63 days when feeding B. tabaci at 25℃, while the shortest development period was found in the group fed on T. urticae at 33℃. Heat stress and B. tabaci significantly decreased A. orientalis fecundity. High temperature significantly reduced the life span of A. orientalis , however, the longevity of A. orientalis feeding on T. urticae were significantly longer than those feeding on B. tabaci , regardless of temperature. Thus, our study assessing A. orientalis performances under different conditions can provide better biological control reference against pest mite and insect by native predatory mites in the fields.
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Both heat stress and prey species affect Amblyseius orientalis performance | 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 Both heat stress and prey species affect Amblyseius orientalis performance Xinyuan Zhou, Hong Yan, Xuemin Hao, Peipei Zhao, Fujing Sheng, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3849776/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 Both temperature and prey closely impact the fitness and predation of natural enemies during biological control. Amblyseius orientalis (Ehara) (Acari: Phytoseiidae) is a native and effective predatory mite against spider mites and small sucking insects in many kinds of orchards and greenhouses, where high temperatures always occur during hot summer. However, the effects of heat stress and prey species on the fitness and predatory efficiency of A. orientalis have not been well understood. In current study, we selected two preys of Tetranychus urticae and Bemisia tabaci and two temperatures of 25℃ and 33℃ to explore the performances of A. orientalis under four combination treatments. The results showed that both temperature and prey significantly affect A. orientalis fitness. In detail, A. orientalis had the longest developmental duration of 7.63 days when feeding B. tabaci at 25℃, while the shortest development period was found in the group fed on T. urticae at 33℃. Heat stress and B. tabaci significantly decreased A. orientalis fecundity. High temperature significantly reduced the life span of A. orientalis , however, the longevity of A. orientalis feeding on T. urticae were significantly longer than those feeding on B. tabaci , regardless of temperature. Thus, our study assessing A. orientalis performances under different conditions can provide better biological control reference against pest mite and insect by native predatory mites in the fields. heat stress prey preference predatory mites spider mites Bemisia tabaci life cycle Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Biological control is a major ecosystem service in agricultural production worldwide, and natural enemies are decisive component in pest management (Martin et al. 2013 ). Many factors can affect biocontrol outcomes, including species identity, resource quality/quantity, and other ecological considerations (Islam et al. 2022 ). Among all biotic and abiotic factors, temperature and prey are primarily important in the ambient environment, since they are directly impact development, reproduction and population establishment of the natural enemy in the ecosystem (Frances and McCauley 2018 ; Bressac et al .2023; Abarca et al . 2021; Price et al. 2011 ). Temperature affects growth, development, physiology, behavior, and geographical distribution of beneficial insects (Yang et al. 2022 ). It can also affect prey-predator foraging interactions by altering their behavioral or physiological responses (Islam et al. 2021 ; Gvoždík et al . 2021). Temperature definitively influences the life traits of insects (Tabebordbar et al. 2022 ; Abou El-Atta et al. 2014 ), for example, the reproduction of Bracon adoxophyesi was significantly affected by high temperature, with longer oviposition period and fewer eggs (Lin et al. 2018 ). In addition, high temperature reducing the longevity and other life traits of natural enemies (Tian et al. 2022 ; Wang et al . 2014). For non-monophagous predators, the type and quantity of nutrients are provided by different preys, which then affect the development and growth of their predators. Harmonia axyridis and Propylaea japonica fed on Corcyra cephalonica could not complete development, but developed well when fed on Myzus persicae (Guo and Wan 2001 ). Prey species also influence their predator’s fecundity. When feeding on pea aphid, Harmonia axyridis had shorter lifespan and lower fecundity than those feeding on Aphis laburni with longer longevity and more eggs (Yu et al. 2018 ). Predatory mites are highly effective natural enemies in controlling pests in agricultural system (Wang et al. 2022 ). Some studies have observed the effects of temperature and prey on the life parameters of predatory mites. For example, high temperatures shortened the development time and the average generation time of Lasioseius japonicus (Zhang et al. 2022 ). The pre-oviposition stage of Neoseiulus womersleyi decreased with the temperature elevation (Sugawara et al. 2017 ). Compared with other prey species, Amblyseius swirskii had the shortest developmental time and the highest reproduction when they fed on Eutetranychus orientalis (Ali and Zaher 2007 ). Amblyseius orientalis (Ehara) (Acari: Phytoseiidae) is a widely distributed predatory mite species in China (Wu and Fang 2021 ). Recent studies have shown that it can prey on both spider mite and other small arthropod pests, such as Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) (Sheng et al. 2014 ; Zhang et al. 2015 ; Yan et al. 2018 ). However, the effects of temperature and different preys on A. orientalis have not been studied. In our study, we selected two preys ( T. urticae and B. tabaci ) and two temperatures (25℃ and 33℃) to explore the performances of A. orientalis under different treatments. Our findings help to understand A. orientalis characteristics interacted by the biotic and abiotic factors, but also provide research basis for better biological control in the fields. Materials and methods Mite rearing and prey culture Amblyseius orientalis was collected from soybean fields in Changli Research Institute of Pomology, Hebei Academy of Agricultural and Forestry Sciences (119°09′E, 39°43′N). The colony had been maintained on Carpoglyphus lactis for 11 years in the Laboratory of Predatory Mites, Institute of Plant Protection, Chinese Academy of Agricultural Sciences (Yan et al. 2022 ). Both mites were reared in an incubator (RXZ, Ningbo, China) at 25 ± 1°C, 70 ± 5% RH and L14:D10 photoperiod (Wei et al. 2023 ). B. tabaci were reared on tobacco, and T. urticae were reared on Phaseolus vulgaris L. Heat tolerance of Amblyseius orientalis To compare the heat tolerance of A. orientalis , T. urticae and B. tabaci , each of 50 one-day old adults were treated by 37℃ and 40℃ for 4 h, respectively. After treatment, all individuals were reared at 25℃, and living numbers were recorded every day for a total of 10 days. Effects of heat and prey on life parameters of Amblyseius orientalis To explore whether high temperature and prey species affect the development of A. orientalis , approximately 100 A. orientalis eggs laid within 12 hours were collected and placed individually in small arenas with sufficient preys of T. urticae or B. tabaci (every stages) at 25℃ and 33℃ respectively. We recorded the time of each developmental stage (egg, larva, protonymph and deutonymph) until adulthood. These adults were then placed in small arenas for mating 24 hours. The gravid females were moved to each arena to observe their reproduction, including the time of pre-oviposition, oviposition, and post-oviposition stages as well as the fecundity and longevity. Effects of heat and prey on predatory capacity of Amblyseius orientalis To examine how high temperature and preys affect predatory efficiency of A. orientalis , 120 one-day old A. orientalis females were collected to starve for 24 h before test. We put each individual in a small arena which had contained 10 eggs of either T. urticae or B. tabaci . They were maintained at 25℃ and 33℃, respectively, and recorded the number of eggs remaining in the arenas after 1 h, 2 h, 4 h and 8 h. 30 replicates for each treatment. Statistical analysis We used Log-rank (Mantel-Cox) test to analyze the survival curves of these three species at two temperatures. Two-way ANOVA was used to examine the effects of temperature and prey on life parameters of A. orientalis . Mann-Whitney U and t test were used to examine impacts of temperature and prey on developmental time, longevity, fecundity and oviposition time. All analyses were run in SPSS 25.0 software and visualized in R 4.3.1. Results Heat tolerance of three species The survival rate of these three species at 37℃ was higher than that at 40℃ (Fig. 1 ) ( A. orientalis : χ (2) 2 = 64.080, P < 0.001 B. tabaci : χ (2) ² = 4.739, P = 0.030; T. urticae : χ (2) ² = 6.585, P = 0.010). No survival difference was found among A. orientalis , B. tabaci , and T. urticae at 37℃ (Fig. 1 A) ( χ (2) ² = 2.498, P = 0.287). Compared with B. tabaci and T. urticae , the survival rate of A. orientalis was significantly lower at 40℃ (Fig. 1 B) ( χ (2) ² = 52.310, P < 0.001). Effects of heat and prey on Amblyseius orientalis development Both temperature and prey affected the developmental duration of A. orientalis (Table 1 ). The developmental time of A. orientalis fed on B. tabaci was significantly longer than that fed on T. urticae . At 25℃, A. orientalis experienced 7.63 days to complete development when feeding on B. tabaci , significantly longer than 6.80 days preyed on T. urticae (Fig. 2 E) (Z = 6.254, P < 0.001). High temperature significantly reduced the developmental period of A. orientalis (Fig. 2 E) ( T. urticae : Z = -10.658, P < 0.001; B. tabaci : Z = -5.384, P < 0.001). Table 1 Effects of temperature and prey on developmental duration, fecundity and longevity of A. orientalis . T: temperature; P: prey; T×P: interaction of temperature and prey. *, P < 0.05; **, P < 0.01; ***, P < 0.001. Traits Term F value p value Egg T F (1,120) = 121 < 0.001*** P F (1,120) = 1.423 0.235 T×P F (1,120) = 6.341 0.013* Larval T F (1,120) = 5.076 0.026* P F (1,120) = 6.341 0.013* T×P F (1,120) = 9.292 0.003** Protonymph T F (1,120) = 28.69 < 0.001*** P F (1,120) = 12.75 0.001** T×P F (1,120) = 11.51 0.001** Deutonymph T F (1,120) = 53.54 < 0.001*** P F (1,120) = 53.54 < 0.001*** T×P F (1,120) = 24.57 < 0.001*** Total developmental time T F (1,120) = 142.4 < 0.001*** P F (1,120) = 53.32 < 0.001*** T×P F (1,120) = 7.123 0.009** Adult longevity T F (1,120) = 82.97 < 0.001*** P F (1,120) = 110.5 < 0.001*** T×P F (1,120) = 18.82 < 0.001*** Total longevity T F (1,120) = 107.3 < 0.001*** P F (1,120) = 95.92 < 0.001*** T×P F (1,120) = 21.74 < 0.001*** Pre-oviposition period T F (1,52) = 2.359 0.131 P F (1,52) = 0.41 0.525 T×P F (1,52) = 2.769 0.102 Oviposition period T F (1,44) = 55.85 < 0.001*** P F (1,44) = 106.3 < 0.001*** T×P F (1,44) = 41.85 < 0.001*** Post-oviposition period T F (1,44) = 40.73 < 0.001*** P F (1,44) = 32.91 < 0.001*** T×P F (1,44) = 36.01 < 0.001*** Lifetime fecundity T F (1,44) = 65.93 < 0.001*** P F (1,44) = 150 < 0.001*** T×P F (1,44) = 47.64 < 0.001*** Daily reproduction T F (1,44) = 3.632 0.063 P F (1,44) = 13.68 0.001** T×P F (1,44) = 3.308 0.076 Effects of heat and prey on fecundity and longevity of Amblyseius orientalis Temperature and prey had no effect on the pre-oviposition stage, but influenced the oviposition period and post-oviposition period of A. orientalis (Table 1 ). The mite had longer oviposition period (Fig. 3 B) (25℃: Z = -7.579, P < 0.001; 33℃: Z = -3.226, P = 0.001) and post-oviposition period (Fig. 3 C) (Z = -6.016, P < 0.001) when fed on T. urticae . The oviposition period and post-oviposition period at 33℃ were shorter than those at 25℃ when they fed on T. urticae (Fig. 3 B & C ) (oviposition: Z = -6.917, P < 0.001; post-oviposition: Z = -5.618, P < 0.001), while the temperature did not work on the oviposition period when mites preying on B. tabaci (Fig. 3 B). Temperature and prey also co-affected fecundity of A. orientalis (Table 1 ). At 25°C, the total egg production of A. orientalis preying on T. urticae was 26.31, which was higher than that preying on B. tabaci (Fig. 3 D) (Z = -7.783, P < 0.001). Whereas, the fecundity decreased significantly at 33℃ regardless of prey species (Fig. 3 D) ( T. urticae : Z = -7.291, P < 0.001; B. tabaci : Z = -1.550, P = 0.121). Fluctuation of daily egg reproduction accounted for prey factor rather than temperature (Table 1 ). Both adult life time and total life time indicated that A. orientalis longevity was significantly longer in the group feeding on T. urticae than on B. tabaci (Fig. 4 ) (25℃: Z = -6.424, P < 0.001; 33℃: Z = -3.805, P < 0.001). High temperature was found to reduce the longevity of A. orientalis with both preys (Fig. 4 , Table 1 ) ( B. tabaci : Z = -3.323, P = 0.001; T. urticae : Z = -6.119, P = 0.014). Effects of heat and prey on predatory capacity of Amblyseius orientalis A. orientalis preferred T. urticae to B. tabaci no matter at 25℃ or 33℃ (Fig. 5 ). Within 8 hours, A. orientalis can prey 7.4 eggs of T. urticae , but only 1.6 eggs of B. tabaci in control condition (Fig. 5 A) (Z = -6.733, P < 0.001,). At 33℃, 6.7 spider mites were preyed by A. orientalis , still higher than 1.5 B. tabaci within 8 hours (Fig. 5 B) (Z = -6.699, P < 0.001). Discussion Widely distributed in China, Amblyseius orientalis can manage spider mites and white flies under the outbreak threshold as an effective natural enemy. Our study indicated that T. urticae and B. tabaci were more heat resistant than A. orientalis , and it’s consistent with previous studies that high temperature was more beneficial to pests than to natural enemies (Stavrinides et al. 2010 ; Stavrinides et al . 2011). High temperature was found to reduce developmental time and fecundity of A. orientalis , confirming the adverse effect of heat stress on predator fitness. Within certain limits, the increase of temperature can accelerate the physiological response and enhance the metabolism (Gillooly et al. 2001 ), leading to the acceleration of development in the organism (Diaz-Cuadros et al. 2023 ). However, high temperature can also accumulate harmful metabolites to damage the organism, resulting in negative life traits and performances (Liu et al. 2022 ). For example, Neoseiulus barkeri prolonged pre-oviposition period and reduced fecundity after thermal stress (Zhang et al. 2016 ). Inappropriate diet can delay insect developmental time (John et al . 1990; Florez-Cuadros et al. 2019 ; Garcia-Robledo et al. 2018 ). Our study confirmed that different preys affected A. orientalis fitness. Food source and nutrient composition from different prey species may account for different fitness of the predator (Ugine et al . 2018; Wen et al. 2020 ). When prey is not suitable, the predator may reduce feeding to cause the delay of growth and development. The limited food intake therefore results in the developmental prolongation and fecundity reduction. For example, the low level of α-linolenic acid and glucose in Corcyra cephalonica affected the metabolic pathway of their predator Arma chinensis to extend developmental period (Li et al. 2016 ). Additionally, we found that both temperature and prey impact on survival and reproduction of A. orientalis . High temperature decreased the developmental period of A. orientalis that fed on spider mites, but not on B. tabaci , indicating that the inappropriate prey had largely affected the predator’s development. It’s not surprising to see that A. orientalis feeding on T. urticae had longer oviposition period and higher fecundity, since spider mite may provide sufficient energy and essential nutrition to maintain long-term reproduction in A. orientalis . It’s consistent with Krol et al. ( 2019 ) that high temperature accelerated the development of mosquito larvae under eutrophic conditions. The biotic factor of diet is more decisive to predator fitness. In sum, we investigated the effects of heat stress and prey species on the survival, longevity, reproduction, and predation of A. orientalis in current study. Our findings showed the sensitive A. orientalis to heat stress and typically negative impacts on fitness. Moreover, B. tabaci was a poor food resource for A. orientalis performances in comparison to T. urticae. Therefore, to achieve a better pest control, the impact of high temperature should be considered when applying A. orientalis in the greenhouse during the hot season. Declarations Acknowledgments This work was supported by the National Key R & D Program of China (2023YFD1400600) and Beijing Innovation Consortium of Agriculture Research System (BAIC01-2024). Author Contributions: Xinyuan Zhou : Formal analysis, Visualization, Writing- Original draft preparation, reviewing and editing. Hong Yan : Writing- Original draft reviewing and editing. Xuemin Hao : Preparation, Investigation. Peipei Zhao : Preparation, Investigation. Fujing Sheng : Preparation, Investigation. Endong Wang : Funding, Supervision and Editing. Xuenong Xu : Funding, Validation, Supervision and Editing. 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Plant protection 44(4):105–9. https://doi.org/10.16688/j.zwbh.2017443 Zhang GH, Li YY, Zhang KJ, Wang JJ, Liu YQ, Liu H (2016) Effects of heat stress on copulation, fecundity and longevity of newly-emerged adults of the predatory mite, Neoseiulus barkeri (Acari: Phytoseiidae). Syst Appl Acarol 21:295. https://doi.org/10.11158/saa.21.3.5 Zhang N, Smith CL, Yin Z, Yan Y, Xie L (2022) Effects of temperature on the adults and progeny of the predaceous mite Lasioseius japonicus (Acari: Blattisociidae) fed on the cereal mite Tyrophagus putrescentiae (Acari: Acaridae). Exp Appl Acarol 86(4):499–515. https://doi.org/10.1007/s10493-022-00708-9 Zhang XX, Lv JL, Hu Y, Wang BM, Chen X, Xu XN, Wang ED (2015) Prey preference and life table of Amblyseius orientalis on Bemisia tabaci and Tetranychus cinnabarinus . PloS one 10(10): e0138820. https://doi.org/10.1371/journal.pone.0138820 Additional Declarations No competing interests reported. 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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-3849776","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":266266440,"identity":"1e286fe4-ec08-4d8d-8eeb-31b8118a470b","order_by":0,"name":"Xinyuan Zhou","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Xinyuan","middleName":"","lastName":"Zhou","suffix":""},{"id":266266441,"identity":"e5d447d0-d136-45d4-b6e5-377ebb7962f5","order_by":1,"name":"Hong Yan","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Hong","middleName":"","lastName":"Yan","suffix":""},{"id":266266442,"identity":"a82fa524-9a36-4477-928d-e297e41fd639","order_by":2,"name":"Xuemin Hao","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Xuemin","middleName":"","lastName":"Hao","suffix":""},{"id":266266443,"identity":"c6518543-b1d7-474a-88ef-2b272f8c8239","order_by":3,"name":"Peipei Zhao","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Peipei","middleName":"","lastName":"Zhao","suffix":""},{"id":266266444,"identity":"8d6b4acc-22c8-4b5a-b8bf-bd4ac4d3d8c2","order_by":4,"name":"Fujing Sheng","email":"","orcid":"","institution":"Shandongzhongke Beneficial Insect Resources Utilization Technology Innovation Center","correspondingAuthor":false,"prefix":"","firstName":"Fujing","middleName":"","lastName":"Sheng","suffix":""},{"id":266266445,"identity":"fb1547ad-e78c-46ab-bd47-2121c5f078fc","order_by":5,"name":"Endong Wang","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Endong","middleName":"","lastName":"Wang","suffix":""},{"id":266266446,"identity":"96f47826-51e9-4d59-9fe6-c42a9d4dab75","order_by":6,"name":"Xuenong Xu","email":"","orcid":"","institution":"Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Xuenong","middleName":"","lastName":"Xu","suffix":""},{"id":266266447,"identity":"bb3a9ea6-e453-45ac-b872-b23299a42602","order_by":7,"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":"2024-01-10 08:58:16","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3849776/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3849776/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":49517993,"identity":"8b2a1d69-ada4-4f83-b7ae-4ac23c7e7419","added_by":"auto","created_at":"2024-01-12 08:54:02","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":127862,"visible":true,"origin":"","legend":"\u003cp\u003eSurvival curves of \u003cem\u003eAmblyseius orientalis\u003c/em\u003e, \u003cem\u003eTetranychus urticae\u003c/em\u003e and \u003cem\u003eBemisia tabaci\u003c/em\u003e after treatments by 37℃ (A) and 40℃ (B). ns, no difference; ***, \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/320866a9022fb52feff88364.png"},{"id":49518384,"identity":"52b3aeee-7379-4094-b149-a673f586e104","added_by":"auto","created_at":"2024-01-12 09:02:02","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":524955,"visible":true,"origin":"","legend":"\u003cp\u003eDevelopmental duration of \u003cem\u003eAmblyseius orientalis \u003c/em\u003efeeding with two preys at two temperatures. (A) Egg stage; (B) Larval stage; (C) Protonymph stage; (D) Deutonymph stage; (E) Total developmental time. The asterisk stands for significant difference. ***, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/dc713b0e9e244cdba86cd0d7.png"},{"id":49517997,"identity":"5a0254a4-9da5-4bdc-9c3c-7d4f3445b5cc","added_by":"auto","created_at":"2024-01-12 08:54:02","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":464519,"visible":true,"origin":"","legend":"\u003cp\u003eReproductive indices of \u003cem\u003eAmblyseius orientalis \u003c/em\u003eunder the different temperatures and preys. (A) Pre-oviposition period; (B) Oviposition period; (C) Post-oviposition period; (D) Lifetime fecundity; (E) Daily reproduction. The asterisks stand for significant difference. ***, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/e36488408232ec18b4a4b6e2.png"},{"id":49517996,"identity":"a8b28e65-682e-494a-bb44-f5b16b127f43","added_by":"auto","created_at":"2024-01-12 08:54:02","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":298263,"visible":true,"origin":"","legend":"\u003cp\u003eThe longevity of \u003cem\u003eA. orientalis\u003c/em\u003e under the different temperature and preys. (A) Adult longevity, (B) Total longevity. *, \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":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/145799e3f5dad5308fe7c892.png"},{"id":49518385,"identity":"77696389-c2dc-418f-a291-cef99460c947","added_by":"auto","created_at":"2024-01-12 09:02:02","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":141113,"visible":true,"origin":"","legend":"\u003cp\u003eConsumed number of \u003cem\u003eT. urticae\u003c/em\u003e and \u003cem\u003eB. tabaci \u003c/em\u003eby \u003cem\u003eA. orientalis\u003c/em\u003e at 25 ℃ (A) and 33 ℃ (B). ***, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.001.\u003c/p\u003e","description":"","filename":"floatimage9.png","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/9ac0ff1c58570ef82c0a8651.png"},{"id":50523895,"identity":"756cf153-c153-4611-92bd-19f53604567f","added_by":"auto","created_at":"2024-02-01 21:07:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1087259,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3849776/v1/95025861-a30d-4c42-9b09-508371fff3bd.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Both heat stress and prey species affect Amblyseius orientalis performance","fulltext":[{"header":"Introduction","content":"\u003cp\u003eBiological control is a major ecosystem service in agricultural production worldwide, and natural enemies are decisive component in pest management (Martin et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Many factors can affect biocontrol outcomes, including species identity, resource quality/quantity, and other ecological considerations (Islam et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Among all biotic and abiotic factors, temperature and prey are primarily important in the ambient environment, since they are directly impact development, reproduction and population establishment of the natural enemy in the ecosystem (Frances and McCauley \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Bressac \u003cem\u003eet al\u003c/em\u003e.2023; Abarca \u003cem\u003eet al\u003c/em\u003e. 2021; Price et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTemperature affects growth, development, physiology, behavior, and geographical distribution of beneficial insects (Yang et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). It can also affect prey-predator foraging interactions by altering their behavioral or physiological responses (Islam et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Gvožd\u0026iacute;k \u003cem\u003eet al\u003c/em\u003e. 2021). Temperature definitively influences the life traits of insects (Tabebordbar et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Abou El-Atta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), for example, the reproduction of \u003cem\u003eBracon adoxophyesi\u003c/em\u003e was significantly affected by high temperature, with longer oviposition period and fewer eggs (Lin et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In addition, high temperature reducing the longevity and other life traits of natural enemies (Tian et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Wang \u003cem\u003eet al\u003c/em\u003e. 2014).\u003c/p\u003e \u003cp\u003eFor non-monophagous predators, the type and quantity of nutrients are provided by different preys, which then affect the development and growth of their predators. \u003cem\u003eHarmonia axyridis\u003c/em\u003e and \u003cem\u003ePropylaea japonica\u003c/em\u003e fed on \u003cem\u003eCorcyra cephalonica\u003c/em\u003e could not complete development, but developed well when fed on \u003cem\u003eMyzus persicae\u003c/em\u003e (Guo and Wan \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Prey species also influence their predator\u0026rsquo;s fecundity. When feeding on pea aphid, \u003cem\u003eHarmonia axyridis\u003c/em\u003e had shorter lifespan and lower fecundity than those feeding on \u003cem\u003eAphis laburni\u003c/em\u003e with longer longevity and more eggs (Yu et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePredatory mites are highly effective natural enemies in controlling pests in agricultural system (Wang et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Some studies have observed the effects of temperature and prey on the life parameters of predatory mites. For example, high temperatures shortened the development time and the average generation time of \u003cem\u003eLasioseius japonicus\u003c/em\u003e (Zhang et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The pre-oviposition stage of \u003cem\u003eNeoseiulus womersleyi\u003c/em\u003e decreased with the temperature elevation (Sugawara et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Compared with other prey species, \u003cem\u003eAmblyseius swirskii\u003c/em\u003e had the shortest developmental time and the highest reproduction when they fed on \u003cem\u003eEutetranychus orientalis\u003c/em\u003e (Ali and Zaher \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAmblyseius orientalis\u003c/em\u003e (Ehara) (Acari: Phytoseiidae) is a widely distributed predatory mite species in China (Wu and Fang \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Recent studies have shown that it can prey on both spider mite and other small arthropod pests, such as \u003cem\u003eBemisia tabaci\u003c/em\u003e (Gennadius) (Hemiptera: Aleyrodidae) (Sheng et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Yan et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, the effects of temperature and different preys on \u003cem\u003eA. orientalis\u003c/em\u003e have not been studied. In our study, we selected two preys (\u003cem\u003eT. urticae\u003c/em\u003e and \u003cem\u003eB. tabaci\u003c/em\u003e) and two temperatures (25℃ and 33℃) to explore the performances of \u003cem\u003eA. orientalis\u003c/em\u003e under different treatments. Our findings help to understand \u003cem\u003eA. orientalis\u003c/em\u003e characteristics interacted by the biotic and abiotic factors, but also provide research basis for better biological control in the fields.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eMite rearing and prey culture\u003c/h2\u003e \u003cp\u003e \u003cem\u003eAmblyseius orientalis\u003c/em\u003e was collected from soybean fields in Changli Research Institute of Pomology, Hebei Academy of Agricultural and Forestry Sciences (119\u0026deg;09\u0026prime;E, 39\u0026deg;43\u0026prime;N). The colony had been maintained on \u003cem\u003eCarpoglyphus lactis\u003c/em\u003e for 11 years in the Laboratory of Predatory Mites, Institute of Plant Protection, Chinese Academy of Agricultural Sciences (Yan et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Both mites were reared in an incubator (RXZ, Ningbo, China) at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and L14:D10 photoperiod (Wei et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u003cem\u003eB. tabaci\u003c/em\u003e were reared on tobacco, and \u003cem\u003eT. urticae\u003c/em\u003e were reared on \u003cem\u003ePhaseolus vulgaris\u003c/em\u003e L.\u003c/p\u003e \u003cp\u003e \u003cb\u003eHeat tolerance of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo compare the heat tolerance of \u003cem\u003eA. orientalis\u003c/em\u003e, \u003cem\u003eT. urticae\u003c/em\u003e and \u003cem\u003eB. tabaci\u003c/em\u003e, each of 50 one-day old adults were treated by 37℃ and 40℃ for 4 h, respectively. After treatment, all individuals were reared at 25℃, and living numbers were recorded every day for a total of 10 days.\u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of heat and prey on life parameters of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo explore whether high temperature and prey species affect the development of \u003cem\u003eA. orientalis\u003c/em\u003e, approximately 100 \u003cem\u003eA. orientalis\u003c/em\u003e eggs laid within 12 hours were collected and placed individually in small arenas with sufficient preys of \u003cem\u003eT. urticae\u003c/em\u003e or \u003cem\u003eB. tabaci\u003c/em\u003e (every stages) at 25℃ and 33℃ respectively. We recorded the time of each developmental stage (egg, larva, protonymph and deutonymph) until adulthood. These adults were then placed in small arenas for mating 24 hours. The gravid females were moved to each arena to observe their reproduction, including the time of pre-oviposition, oviposition, and post-oviposition stages as well as the fecundity and longevity.\u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of heat and prey on predatory capacity of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTo examine how high temperature and preys affect predatory efficiency of \u003cem\u003eA. orientalis\u003c/em\u003e, 120 one-day old \u003cem\u003eA. orientalis\u003c/em\u003e females were collected to starve for 24 h before test. We put each individual in a small arena which had contained 10 eggs of either \u003cem\u003eT. urticae\u003c/em\u003e or \u003cem\u003eB. tabaci\u003c/em\u003e. They were maintained at 25℃ and 33℃, respectively, and recorded the number of eggs remaining in the arenas after 1 h, 2 h, 4 h and 8 h. 30 replicates for each treatment.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eWe used Log-rank (Mantel-Cox) test to analyze the survival curves of these three species at two temperatures. Two-way ANOVA was used to examine the effects of temperature and prey on life parameters of \u003cem\u003eA. orientalis\u003c/em\u003e. Mann-Whitney U and t test were used to examine impacts of temperature and prey on developmental time, longevity, fecundity and oviposition time. All analyses were run in SPSS 25.0 software and visualized in R 4.3.1.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eHeat tolerance of three species\u003c/h2\u003e \u003cp\u003eThe survival rate of these three species at 37℃ was higher than that at 40℃ (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) (\u003cem\u003eA. orientalis\u003c/em\u003e: \u003cem\u003eχ\u003c/em\u003e\u003csub\u003e(2)\u003c/sub\u003e\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;64.080, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001 \u003cem\u003eB. tabaci\u003c/em\u003e: \u003cem\u003eχ\u003c/em\u003e\u003csub\u003e(2)\u003c/sub\u003e\u0026sup2; = 4.739, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.030; \u003cem\u003eT. urticae\u003c/em\u003e: \u003cem\u003eχ\u003c/em\u003e\u003csub\u003e(2)\u003c/sub\u003e\u0026sup2; = 6.585, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.010). No survival difference was found among \u003cem\u003eA. orientalis\u003c/em\u003e, \u003cem\u003eB. tabaci\u003c/em\u003e, and \u003cem\u003eT. urticae\u003c/em\u003e at 37℃ (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA) (\u003cem\u003eχ\u003c/em\u003e\u003csub\u003e(2)\u003c/sub\u003e\u0026sup2; = 2.498, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.287). Compared with \u003cem\u003eB. tabaci\u003c/em\u003e and \u003cem\u003eT. urticae\u003c/em\u003e, the survival rate of \u003cem\u003eA. orientalis\u003c/em\u003e was significantly lower at 40℃ (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB) (\u003cem\u003eχ\u003c/em\u003e\u003csub\u003e(2)\u003c/sub\u003e\u0026sup2; = 52.310, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of heat and prey on\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e \u003cb\u003edevelopment\u003c/b\u003e\u003c/p\u003e \u003cp\u003eBoth temperature and prey affected the developmental duration of \u003cem\u003eA. orientalis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The developmental time of \u003cem\u003eA. orientalis\u003c/em\u003e fed on \u003cem\u003eB. tabaci\u003c/em\u003e was significantly longer than that fed on \u003cem\u003eT. urticae\u003c/em\u003e. At 25℃, \u003cem\u003eA. orientalis\u003c/em\u003e experienced 7.63 days to complete development when feeding on \u003cem\u003eB. tabaci\u003c/em\u003e, significantly longer than 6.80 days preyed on \u003cem\u003eT. urticae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eE) (Z\u0026thinsp;=\u0026thinsp;6.254, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). High temperature significantly reduced the developmental period of \u003cem\u003eA. orientalis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eE) (\u003cem\u003eT. urticae\u003c/em\u003e: Z = -10.658, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; \u003cem\u003eB. tabaci\u003c/em\u003e: Z = -5.384, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\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\u003eEffects of temperature and prey on developmental duration, fecundity and longevity of \u003cem\u003eA. orientalis\u003c/em\u003e. T: temperature; P: prey; T\u0026times;P: interaction of temperature and prey. *, \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\u003eF 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\u003eEgg\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;121\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1.423\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.235\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;6.341\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.013*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eLarval\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;5.076\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.026*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;6.341\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.013*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;9.292\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.003**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eProtonymph\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;28.69\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;12.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;11.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eDeutonymph\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;53.54\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;53.54\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;24.57\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\u003eTotal developmental time\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;142.4\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;53.32\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;7.123\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.009**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eAdult longevity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;82.97\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;110.5\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;18.82\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\u003eTotal longevity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;107.3\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;95.92\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,120)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;21.74\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\u003ePre-oviposition period\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,52)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;2.359\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.131\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,52)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.525\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,52)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;2.769\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.102\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eOviposition period\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;55.85\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;106.3\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;41.85\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\u003ePost-oviposition period\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;40.73\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;32.91\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;36.01\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\u003eLifetime fecundity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;65.93\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\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;150\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\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;47.64\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\u003eDaily reproduction\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;3.632\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.063\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;13.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eT\u0026times;P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u003csub\u003e(1,44)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;3.308\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.076\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\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of heat and prey on fecundity and longevity of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003eTemperature and prey had no effect on the pre-oviposition stage, but influenced the oviposition period and post-oviposition period of \u003cem\u003eA. orientalis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The mite had longer oviposition period (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB) (25℃: Z = -7.579, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; 33℃: Z = -3.226, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001) and post-oviposition period (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC) (Z = -6.016, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) when fed on \u003cem\u003eT. urticae\u003c/em\u003e. The oviposition period and post-oviposition period at 33℃ were shorter than those at 25℃ when they fed on \u003cem\u003eT. urticae\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB \u0026amp; C ) (oviposition: Z = -6.917, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; post-oviposition: Z = -5.618, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), while the temperature did not work on the oviposition period when mites preying on \u003cem\u003eB. tabaci\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eTemperature and prey also co-affected fecundity of \u003cem\u003eA. orientalis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). At 25\u0026deg;C, the total egg production of \u003cem\u003eA. orientalis\u003c/em\u003e preying on \u003cem\u003eT. urticae\u003c/em\u003e was 26.31, which was higher than that preying on \u003cem\u003eB. tabaci\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eD) (Z = -7.783, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Whereas, the fecundity decreased significantly at 33℃ regardless of prey species (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eD) (\u003cem\u003eT. urticae\u003c/em\u003e: Z = -7.291, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; \u003cem\u003eB. tabaci\u003c/em\u003e: Z = -1.550, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.121). Fluctuation of daily egg reproduction accounted for prey factor rather than temperature (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBoth adult life time and total life time indicated that \u003cem\u003eA. orientalis\u003c/em\u003e longevity was significantly longer in the group feeding on \u003cem\u003eT. urticae\u003c/em\u003e than on \u003cem\u003eB. tabaci\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e) (25℃: Z = -6.424, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; 33℃: Z = -3.805, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). High temperature was found to reduce the longevity of \u003cem\u003eA. orientalis\u003c/em\u003e with both preys (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) (\u003cem\u003eB. tabaci\u003c/em\u003e: Z = -3.323, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001; \u003cem\u003eT. urticae\u003c/em\u003e: Z = -6.119, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.014).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eEffects of heat and prey on predatory capacity of\u003c/b\u003e \u003cb\u003eAmblyseius orientalis\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003cem\u003eA. orientalis\u003c/em\u003e preferred \u003cem\u003eT. urticae\u003c/em\u003e to \u003cem\u003eB. tabaci\u003c/em\u003e no matter at 25℃ or 33℃ (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Within 8 hours, \u003cem\u003eA. orientalis\u003c/em\u003e can prey 7.4 eggs of \u003cem\u003eT. urticae\u003c/em\u003e, but only 1.6 eggs of \u003cem\u003eB. tabaci\u003c/em\u003e in control condition (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eA) (Z = -6.733, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001,). At 33℃, 6.7 spider mites were preyed by \u003cem\u003eA. orientalis\u003c/em\u003e, still higher than 1.5 \u003cem\u003eB. tabaci\u003c/em\u003e within 8 hours (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eB) (Z = -6.699, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eWidely distributed in China, \u003cem\u003eAmblyseius orientalis\u003c/em\u003e can manage spider mites and white flies under the outbreak threshold as an effective natural enemy. Our study indicated that \u003cem\u003eT. urticae\u003c/em\u003e and \u003cem\u003eB. tabaci\u003c/em\u003e were more heat resistant than \u003cem\u003eA. orientalis\u003c/em\u003e, and it\u0026rsquo;s consistent with previous studies that high temperature was more beneficial to pests than to natural enemies (Stavrinides et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Stavrinides \u003cem\u003eet al\u003c/em\u003e. 2011). High temperature was found to reduce developmental time and fecundity of \u003cem\u003eA. orientalis\u003c/em\u003e, confirming the adverse effect of heat stress on predator fitness. Within certain limits, the increase of temperature can accelerate the physiological response and enhance the metabolism (Gillooly et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), leading to the acceleration of development in the organism (Diaz-Cuadros et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, high temperature can also accumulate harmful metabolites to damage the organism, resulting in negative life traits and performances (Liu et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). For example, \u003cem\u003eNeoseiulus barkeri\u003c/em\u003e prolonged pre-oviposition period and reduced fecundity after thermal stress (Zhang et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInappropriate diet can delay insect developmental time (John \u003cem\u003eet al\u003c/em\u003e. 1990; Florez-Cuadros et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Garcia-Robledo et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Our study confirmed that different preys affected \u003cem\u003eA. orientalis\u003c/em\u003e fitness. Food source and nutrient composition from different prey species may account for different fitness of the predator (Ugine \u003cem\u003eet al\u003c/em\u003e. 2018; Wen et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). When prey is not suitable, the predator may reduce feeding to cause the delay of growth and development. The limited food intake therefore results in the developmental prolongation and fecundity reduction. For example, the low level of α-linolenic acid and glucose in \u003cem\u003eCorcyra cephalonica\u003c/em\u003e affected the metabolic pathway of their predator \u003cem\u003eArma chinensis\u003c/em\u003e to extend developmental period (Li et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAdditionally, we found that both temperature and prey impact on survival and reproduction of \u003cem\u003eA. orientalis\u003c/em\u003e. High temperature decreased the developmental period of \u003cem\u003eA. orientalis\u003c/em\u003e that fed on spider mites, but not on \u003cem\u003eB. tabaci\u003c/em\u003e, indicating that the inappropriate prey had largely affected the predator\u0026rsquo;s development. It\u0026rsquo;s not surprising to see that \u003cem\u003eA. orientalis\u003c/em\u003e feeding on \u003cem\u003eT. urticae\u003c/em\u003e had longer oviposition period and higher fecundity, since spider mite may provide sufficient energy and essential nutrition to maintain long-term reproduction in \u003cem\u003eA. orientalis\u003c/em\u003e. It\u0026rsquo;s consistent with Krol et al. (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) that high temperature accelerated the development of mosquito larvae under eutrophic conditions. The biotic factor of diet is more decisive to predator fitness.\u003c/p\u003e \u003cp\u003eIn sum, we investigated the effects of heat stress and prey species on the survival, longevity, reproduction, and predation of \u003cem\u003eA. orientalis\u003c/em\u003e in current study. Our findings showed the sensitive \u003cem\u003eA. orientalis\u003c/em\u003e to heat stress and typically negative impacts on fitness. Moreover, \u003cem\u003eB. tabaci\u003c/em\u003e was a poor food resource for \u003cem\u003eA. orientalis\u003c/em\u003e performances in comparison to \u003cem\u003eT. urticae.\u003c/em\u003e Therefore, to achieve a better pest control, the impact of high temperature should be considered when applying \u003cem\u003eA. orientalis\u003c/em\u003e in the greenhouse during the hot season.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by the National Key R \u0026amp; D Program of China (2023YFD1400600) and Beijing Innovation Consortium of Agriculture Research System (BAIC01-2024).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAuthor Contributions:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eXinyuan Zhou\u003c/strong\u003e: Formal analysis, Visualization, Writing- Original draft preparation, reviewing and editing. \u003cstrong\u003eHong Yan\u003c/strong\u003e: Writing- Original draft reviewing and editing. \u003cstrong\u003eXuemin Hao\u003c/strong\u003e: Preparation, Investigation. \u003cstrong\u003ePeipei Zhao\u003c/strong\u003e: Preparation, Investigation. \u003cstrong\u003eFujing Sheng\u003c/strong\u003e: Preparation, Investigation. \u003cstrong\u003eEndong Wang\u003c/strong\u003e: Funding, Supervision and Editing. \u003cstrong\u003eXuenong Xu\u003c/strong\u003e: Funding, Validation, Supervision and Editing. \u003cstrong\u003eBo Zhang\u003c/strong\u003e: Conception, Funding, Supervision and Writing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have declared that no competing interest exits.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbarca M, Spahn R (2021) Direct and indirect effects of altered temperature regimes and phenological mismatches on insect populations.\u003cem\u003e \u003c/em\u003eCurr Opin Insect Sci 47:67\u0026ndash;74. https://doi.org/10.1016/j.cois.2021.04.008\u003c/li\u003e\n\u003cli\u003eAbou El-Atta DA, Ghazy NA, Osman MA (2014) Effects of temperature on the life-history traits of \u003cem\u003eSancassania (Caloglyphus) berlesei\u003c/em\u003e (Acari: Astigmatina: Acaridae) feeding on root-knot nematodes, \u003cem\u003eMeloidogyne\u003c/em\u003e spp. (Nematoda: Meloidogynidae). 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PloS one 10(10): e0138820. https://doi.org/10.1371/journal.pone.0138820\u003c/li\u003e\n\u003c/ol\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":"heat stress, prey preference, predatory mites, spider mites, Bemisia tabaci, life cycle","lastPublishedDoi":"10.21203/rs.3.rs-3849776/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3849776/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eBoth temperature and prey closely impact the fitness and predation of natural enemies during biological control. \u003cem\u003eAmblyseius orientalis\u003c/em\u003e (Ehara) (Acari: Phytoseiidae) is a native and effective predatory mite against spider mites and small sucking insects in many kinds of orchards and greenhouses, where high temperatures always occur during hot summer. However, the effects of heat stress and prey species on the fitness and predatory efficiency of \u003cem\u003eA. orientalis\u003c/em\u003e have not been well understood. In current study, we selected two preys of \u003cem\u003eTetranychus urticae\u003c/em\u003e and \u003cem\u003eBemisia tabaci\u003c/em\u003e and two temperatures of 25℃ and 33℃ to explore the performances of \u003cem\u003eA. orientalis\u003c/em\u003e under four combination treatments. The results showed that both temperature and prey significantly affect \u003cem\u003eA. orientalis\u003c/em\u003e fitness. In detail, \u003cem\u003eA. orientalis\u003c/em\u003e had the longest developmental duration of 7.63 days when feeding \u003cem\u003eB. tabaci\u003c/em\u003e at 25℃, while the shortest development period was found in the group fed on \u003cem\u003eT. urticae\u003c/em\u003e at 33℃. Heat stress and \u003cem\u003eB. tabaci\u003c/em\u003e significantly decreased \u003cem\u003eA. orientalis\u003c/em\u003e fecundity. High temperature significantly reduced the life span of \u003cem\u003eA. orientalis\u003c/em\u003e, however, the longevity of \u003cem\u003eA. orientalis\u003c/em\u003e feeding on \u003cem\u003eT. urticae\u003c/em\u003e were significantly longer than those feeding on \u003cem\u003eB. tabaci\u003c/em\u003e, regardless of temperature. Thus, our study assessing \u003cem\u003eA. orientalis\u003c/em\u003e performances under different conditions can provide better biological control reference against pest mite and insect by native predatory mites in the fields.\u003c/p\u003e","manuscriptTitle":"Both heat stress and prey species affect Amblyseius orientalis performance","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-01-12 08:53:58","doi":"10.21203/rs.3.rs-3849776/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":"January 12th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-02-01T20:59:49+00:00","versionOfRecord":[],"versionCreatedAt":"2024-01-12 08:53:58","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3849776","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3849776","identity":"rs-3849776","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}

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