Comparison of the predatory impacts of indigenous and adventive ladybeetle species (Coleoptera: Coccinellidae) using a functional response approach

Comparison of the predatory impacts of indigenous and adventive ladybeetle species (Coleoptera: Coccinellidae) using a functional response approach | 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 Comparison of the predatory impacts of indigenous and adventive ladybeetle species (Coleoptera: Coccinellidae) using a functional response approach Fateme Ranjbar, M. Amin Jalali, Zahra Ahmadi, Xavier Pons, Alexandre Levi-Mourao, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3871729/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 23 May, 2024 Read the published version in Journal of Plant Diseases and Protection → Version 1 posted 4 You are reading this latest preprint version Abstract Biological control has long been recognized for its economic and ecological benefits as a pest management approach, but the use of non-native natural enemies has raised concerns about potential risks and unintended consequences. Indigenous predatory ladybeetles, such as Oenopia conglobata , play a crucial role in controlling many sap-sucking pests, including Agonoscena pistaciae , which is the most destructive pest of pistachio trees in Iran and other pistachio-growing regions worldwide. However, the abundance of Oenopia conglobata has recently decreased due to the presence of an adventive predatory ladybeetle, Menochilus sexmaculatus . To better understand the potential risks associated with this adventive species, we investigated the functional responses of female adults of M. sexmaculatus and O. conglobata against the third and fourth instar nymphs of A. pistaciae and Aphis gossypii . Our findings revealed that both predators exhibited a type II functional response, with O. conglobata demonstrating a significantly higher attack rate against A. pistaciae nymphs than A. gossypii nymphs. In contrast, prey species did not have a significant effect on the attack rate of M. sexmaculatus . Notably, M. sexmaculatus displayed the highest predation rate and voracity against both prey species. These results provide valuable insights into the potential risks of M. sexmaculatus for indigenous predatory species like O. conglobata . The fact that M. sexmaculatus exhibits higher voracity for both prey species than O. conglobata does suggest that it may pose a threat to the native ladybeetle population, highlighting the need for further research and careful consideration when introducing non-native natural enemies for pest management. Agonoscena pistaciae Aphis gossypii prey density Oenopia conglobata Menochilus sexmaculatus Figures Figure 1 Figure 2 Introductions Pistachio is one of the most economically important crops in Iran, covering ~ 420,000 ha of land (Mehrnejad, 2010 ). The common pistachio psyllid – Agonoscena pistaciae Burckhardt and Lauterer (Hemiptera: Psyllidae) – poses a significant threat to cultivated pistachio trees in the Russian Federation, Middle East and Mediterranean areas (Mehrnejad et al. 2010; 2015 ). This insect has also evolved resistance to diverse pesticides, making chemical control strategies less effective (Mehrnejad, 2010 ). Biological control agents offer a promising alternative as part of integrated pest management systems, particularly as a strategy to combat insecticide-resistant pests (Bale et al. 2008). Coccinellids (Col. Coccinellidae), also known as ladybeetles, ladybugs or ladybirds, are beneficial insects in agricultural ecosystems because they help to maintain equilibrium and biological control for a wide range of pests (Obrycki and Kring 1998 ; Biddinger et al., 2009 ; Kundoo and Khan 2017 ). Several predatory ladybeetle species are found in Rafsanjan, the most important pistachio-growing region in Iran (Mehrnejad, 2010 ). Predatory ladybeetles, along with other biological control agents in the area, control the population of the common pistachio psyllid, which increases rapidly from the emergence of shoots in early spring to early summer and remains stable on pistachio trees until the autumn leaf drop. Nymphs and adult stages of the common pistachio psyllid feed on sap leaves, excreting significant amounts of honeydew that, when dry, lead to leaf and fruit scorching. This feeding behavior has severe economic consequences, such as reduces plant vigor, causing leaf drop, stunting, low fruit yields and the shedding of shoots (Mehrnejad, 2001 ). During spring, aphids such as Aphis gossypii Glover (cotton aphid) and Aphis craccivora Koch (cowpea aphid) become established on herbaceous plants earlier than the common pistachio psyllid (Jalali, 2001 ). Adult ladybeetles gradually emerge from their overwintering sites and disperse on the plants at the edges of pistachio orchards. They engage in mating and egg-laying while also feeding heavily on aphids. At this time, the pistachio tree buds have not yet opened, but the adult psyllids, which typically appear from mid-February onwards, have already infested the pistachio branches. As the common pistachio psyllid begins egg-laying on the swollen buds of pistachio trees, the adult coccinellids begin to feed on these pests. Oenopia conglobata (Linnaeus) is the most abundant ladybeetle in pistachio orchards. It shows greater activity on pistachio trees even when understory plants such as Alhagi camelorum are infested with aphids. Moreover, when the pistachio psyllid population is not available in sufficient numbers on pistachio trees, O. conglobata feeds on aphids infesting understory plants (Jalali, 2001 ; Mehrnejad and Jalali 2004 ). Menochilus sexmaculatus (Fabricius) is another important predator of aphids, psyllids, thrips, whiteflies, moth larvae, and mites (Sugiura, 1998 ; Saleem et al., 2014 ). Both predators are important biological control agents in pistachio orchards, but O. conglobata is indigenous whereas M. sexmaculatus is recently adventive (Jalali, 2001 ; Ranjbar et al., 2020 ). Each species readily feeds on A . pistaciae and A. gossypii (Mehrnejad and Jalali, 2004 ). One of the most important behavioral characteristics of natural enemies from a pest management perspective is the attack rate ( a ) against prey (Holling, 1966 ). However, this is limited by the time needed by the predator to subdue and consume prey, also known as the handling time ( T h ). Both parameters determine the functional response of a natural enemy against prey (Holling, 1966 ). This predicts how the predatory behavior of a natural enemy will develop when the prey density increases, and can be used to determine the potential of natural enemies as biological control agents. Three types of functional responses were first described in the mid-twentieth century (Solomon 1949 ). In a type I response, the predator’s consumption rate increases in linearly with the prey density up to the satiation point, whereupon consumption levels plateau and stabilize. In a type II response, the predation rate declines and approaches an asymptote as prey density increases. This occurs because predators are constrained by the time required to handle prey. Finally, a type III response is characterized by an increasing predation rate that correlates positively with prey density. However, there is an inflection point, resulting in an asymptote, when the prey density surpasses the capacity to handle additional prey (Holling 1959a ). When assessing functional responses, it should be noted that the type of response and its parameters can be influenced by factors such as temperature, host plant, prey life stage, the age, sex and developmental stage of the predator, and the spatial scale (McCaffrey and Horsburgh 1986 ; Farhadi et al., 2010 ; Jalali et al., 2010 ; Madadi et al., 2012; Jalali and Ziaaddini 2017 ; Uiterwaal and DeLong 2018; Sanati et al., 2020 ; Yuliadhi et al., 2021 ; Juliano et al., 2022 ). Despite the important role of O. conglobata and M. sexmaculatus in pistachio orchards, their functional responses to the common pistachio psyllid population are unknown. We therefore investigated the functional responses of both ladybeetle species against the common pistachio psyllid and the cotton aphid. This will help to determine the value of these ladybeetle species in biological control strategies, taking into account any potential interference caused by the cooccurrence of the two pests. Our data provide valuable insights into the effectiveness and behavior of these ladybeetle species as biological control agents in pistachio orchards, and can be used in the future to optimize pest management strategies. Material and Methods Insect colonies Predatory coccinellids M. sexmaculatus and O. conglobata colonies were established from several collections of adults made in pistachio orchards of Rafsanjan during the spring season 2022. Individuals were found on pistachio trees infested with the common pistachio psyllid or on understory plants feeding on aphids such as A. gossypii and A. craccivora . The orchards selected in collection sites had not been treated with pesticides. The ladybeetles were transported in plastic containers with suitable ventilation to an insect rearing room (26 ± 1°C, 65% ± 5% relative humidity, 16:8 L:D photoperiod). After sorting and identifying the species, adult specimens were transferred to larger plastic containers (23 × 18 × 8 cm) with appropriate ventilation. To increase the population, male and female mating pairs were placed in 12-cm Petri dishes lined with paper towels and vented with mesh-covered holes in the lid. To minimize any potential effect of conditioning caused by the feeding on a specific type of prey, each ladybeetle species was reared on a diet of Mediterranean flour moth ( Ephestia kuehniella Zeller) eggs for five generations prior to their use in the experiments. For the mating pairs in Petri dishes, we provided two small plastic containers, one filled with moistened cotton wool to provide humidity and the other containing ~ 0.1 g fresh weight of flour moth eggs, which were checked daily (Ranjbar et al., 2020 ). If ladybeetle eggs were found when the Petri dishes were checked, the adults were transferred to new dishes and the containers with the eggs were maintained until the eggs hatched. This management approach allowed for the continuous rearing of ladybeetle populations. Aphis gossypii The initial population of cotton aphids was collected from infested greenhouses. To establish a colony, squash plants ( Cucurbita pepo L.) were used as hosts. Seeds were sown in plastic pots (12 × 12.5 cm) containing a 1:1 mixture of cocopeat and perlite before transfer to a growth chamber. The plants were then transferred to mesh cages, which were used to establish a colony of aphids. Agonoscena pistaciae Fresh pistachio leaves infested with common pistachio psyllid were collected from the experimental pistachio garden of Vali-e-Asr University of Rafsanjan, Iran. These insects were used for the functional response experiments within 12 h of collection. Functional response test system Functional responses were tested by exposing mated female M. sexmaculatus or O. conglobata (< 7 days old, in the process of egg laying) to prey densities of 10, 20, 40, 60, 80, 160 and 200 third and fourth instar nymphs of A. gossypii or third and fourth instar A. pistaciae under laboratory conditions (26 ± 1°C temperature, 65% ± 5% relative humidity, 16:8 L:D photoperiod). To standardize the experimental conditions, ladybeetles were starved for 24 h before starting the experiments. We then transferred the nymphs to 12-cm Petri dishes containing a fully mature pistachio leaf (for psyllids) or a 4-day-old cucumber leaf (for aphids). The leaf petiole was placed within an Eppendorf tube filled with water to preserve leaf turgor. Dishes were prepared at different prey densities (For each density, there were 10 biological replicates conducted over the course of a week, with all densities included in one replicate on a single day.) and we counted the number of prey remaining after 24 h. Functional response data analysis The type of functional response was determined by logistic regression, with the ratio of prey consumed as a function of the initial prey density, described by a multinomial function according to Eq. 1 (De Clercq et al. 2000 , Juliano 2001 ): where N e is the number of prey consumed, N 0 is the initial prey density, and P 0 – P 3 are parameters that need to be estimated. The sign (negative or positive) of the linear coefficient in the polynomial function indicates the type of second-order and third-order functional responses, respectively (Juliano, 2001 ). The data for the functional response were used as inputs for the random predator equation (Rogers,1972; Juliano, 2001 ). Equations 2 and 3 correspond to the type II and type III functional responses, respectively: where N e is the number of preys eaten, N 0 is the initial prey density, a is the attack constant, T is the total time available (24 h), T h is the handling time per prey, and b , c and d are constants. By applying asymptotic 95% confidence intervals (95% CI), we could statistically separate the parameter estimates (Juliano 2001 ). Analysis of variance (ANOVA) was applied using SPSS for variance analysis and SAS (SAS Institute 1989) for functional response data analysis. One-way ANOVA was used to compare the functional response parameters. Results A negative correlation coefficient was revealed by linear regression analysis (P1 < 0) for both ladybeetle species, indicating a type II functional response when feeding on third and fourth instar nymphs of A. gossypii and A. pistaciae at different densities (Table 1 , Fig. 1 ). The logistic regression of prey consumed by female ladybeetles at different initial prey densities indicated the significance of the linear portion of the regression equation (P < 0.05). The decline in the number of prey consumed by ladybeetles with increasing prey density revealed the non-linearity of the regression equation (Table 1 ). The attack rate of O. conglobata was significantly higher against nymphs of A. pistaciae compared to A. gossypii . However, the attack rate of M. sexmaculatus did not differ significantly between the prey species (Table 2 ). The prey handling time for both ladybeetle species was higher when feeding on A. gossypii compared to A. pistaciae (Table 2 , Fig. 2 ). In a 1-h time interval, both ladybeetle species captured a greater number ( T / T h ) of A. pistaceae compared to A. gossypii (Table 2 ). Table 1 Results of logistic regression analysis of the proportion of Agonoscena pistaciae or Aphis gossypii nymphs (3rd and 4th instars) eaten by adult females of two predatory ladybeetles, Oenopia conglobata and Menochilus sexmaculatus , on initial prey density. Predator Prey Coefficient Estimate SE Chi Squqre P-value A. pistaciae Constant ( P 0 ) 4.4147 0.4791 84.8955 < .0001 Linear ( P 1 ) -0.0418 0.0133 9.8208 0.0017 O. conglobata Quadratic ( P 2 ) 0.000014 0.000109 0.0173 0.8955 Cubic ( Pc 3 ) 3.631E-7 2.768E-7 1.7204 0.1896 A. gossypii Constant ( P 0 ) 2.2662 0.2600 75.9737 < .0001 Linear ( P 1 ) -0.0139 0.00799 3.0111 0.0827 Quadratic ( P 2 ) -0.00009 0.000071 1.6465 0.1994 Cubic ( Pc 3 ) 4.321E-7 1.944E-7 4.9411 0.0262 A. pistaciae Constant ( P 0 ) 5.7194 0.6261 83.4486 < .0001 Linear ( P 1 ) -0.0748 0.0169 19.6006 < .0001 M. sexmaculatus Quadratic ( P 2 ) 0.000297 0.000135 4.8586 0.0275 Cubic ( Pc 3 ) -3.79E-7 3.324E-7 1.3021 0.2538 A. gossypii Constant ( P 0 ) 2.5616 0.3033 71.3431 < .0001 Linear ( P 1 ) -0.00559 0.00915 0.3733 0.5412 Quadratic ( P 2 ) -0.00018 0.000080 5.1177 0.0237 Cubic ( Pc 3 ) 6.604E-7 2.124E-7 9.6705 0.0019 Table 2 Parameters estimated by the random predator equation and corresponding asymptotic 95% confidence intervals (CI) indicating functional response of adult females of Oenopia conglobata or Menochilus sexmaculatus to densities of 3rd -4th instars of two natural preys, Agonoscena pistaciae and Aphis gossypii . Asymptotic 95% CI Asymptotic 95% CI Predator Prey Type a † Lower Upper T h ‡ Lower Upper T/ T h * r 2 § O. conglobata A. pistaciae II 0.315 ± 0.016 a 0.280 0.350 0.317 ± 0.001 b 0.315 0.319 75 0.962 A. gossypii II 0.172 ± 0.006 c 0.159 0.185 0.340 ± 0.003 a 0.333 0.347 70 0.961 M. sexmaculatus A. pistaciae II 0.274 ± 0.016 ab 0.238 0.310 0.287 ± 0.001 d 0.283 0.290 83 0.985 A. gossypii II 0.229 ± 0.01 b 0.207 0.251 0.296 ± 0.002 c 0.291 0.301 81 0.981 F = 21.554 F = 123.263 df = 3,48 df = 3,48 P < 0.001 P < 0.001 † attack rate expressed as prey consumed per hour. ‡ T h handling time expressed as hour(s). * T/ T h estimated maximum numbers of 3rd and 4th instar nymphs of Agonoscena pistaciae or Aphis gossypii , preyed by one Oenopia conglobata or Menochilus sexmaculatus per hour. § The coefficient of determination [1- (residual sum of squares/corrected total sum of squares)]. Means (± SE) within the same column followed by the same letter are statistically equivalent (Tukey–Kramer HSD test, α = 0.05). Discussion We used logistic regression analysis to determine the type of functional response exhibited by females of the ladybeetle species O. conglobata and M . sexmaculatus when feeding on the third and fourth instar nymphs of A. gossypii and A . pistaciae . Our calculations revealed a type II response for both predators against both prey types, indicating that the more the prey density increases, the lower the proportion of prey captured. In other words, the predator’s response is density dependent and inversely correlated with the number of prey. At low prey densities, the predator spends more time searching for prey, whereas at high prey densities it invests more time in capturing the prey (Holling, 1959b ). In many previous studies, a type II functional response has been documented for various species of predatory ladybeetles when interacting with different prey under diverse laboratory conditions (Hasani et al., 2008 ; Khan and Mir, 2008 ; Jafari and Goldasteh 2009 ; Sabaghi et al., 2011 ; Madadi et al., 2011 ; Yuliadhi et al., 2021 ; Pervez et al., 2022 ). In some cases, the abundance of various life stages of predatory ladybeetles was positively correlated with prey densities (Hodek, 1973 ; Agarwala & Bardhanroy, 1997 ; Dixon 1997 ; Koch, 2003 ; Rana, 2006 ; Meseguer et al. 2021 ). Therefore, some researchers have questioned the efficiency of predatory insects with type II functional responses as biological control agents. Although the shape of the functional response curve is important, it may not be sufficient as a sole criterion for predicting the success or failure of a predator in a biological control program. Other factors are also relevant, such as the numerical response, the intrinsic rate of population increase, and the competitive ability of the predator (Pervez and Omkar, 2005 ). However, some predators with a type II functional response in laboratory experiments have nevertheless achieved success in mass release programs against pests (Foglar et al., 1990 ; Tedders and Schaefer, 1994 ). In assessing the potential effectiveness of natural enemies, parameters such as the handling time ( T h ) and attack rate ( a ) should be taken into account. We found that the attack rate of O. conglobata against the common pistachio psyllid was significantly higher than against the cotton aphid. In contrast, M. sexmaculatus attacked each prey species at a similar rate. The longest handling times were recorded for O. conglobata feeding on cotton aphids, followed by the same predator feeding on the common pistachio psyllid, and then M. sexmaculatus feeding on the cotton aphid, and finally M. sexmaculatus feeding on the common pistachio psyllid. The two predator species, M. sexmaculatus and O. conglobata , exhibited a higher consumption rate of A. gossypii compared to A. pistaciae . Moreover, M . sexmaculatus demonstrated greater predation on both prey species in comparison to O . conglobata . The mean fresh weight of a mixture containing 500 third and fourth instar nymphs of A. gossypii and A. pistaciae (n = ten replicates of 50 nymphs each) was estimated to be 0.283 ± 0.009 mg and 0.261 ± 0.010 mg, respectively (Jalali et al., 2018 ). Despite the equivalent biomass of third and fourth instars in both prey species, this result is likely influenced by the larger size of M. sexmaculatus , which may be associated with higher voracity or variations in nutritional indices of the prey. It appears that aphid individuals contain more energy than psyllids (Ranjbar et al., 2020 ).An increase in handling time reduced the number of aphids or psyllids consumed by a single predator individual in 1 h. The handling time for both predators feeding on the cotton aphid was significantly longer compared to their feeding on the common pistachio psyllid. Therefore, they consumed more psyllids than aphids. The fourth instar larvae and adult ladybeetles are considered the key developmental stages for biological control due to their high feeding capacity (Hodek and Honek, 1996 ). Given that the release of fourth-instar larvae is challenging under natural conditions due to their vulnerability, adult ladybeetles are more suitable for release. The predation rate of female insects is generally higher than that of males (Ghosh and Agarwala 2018 ; Islam et al., 2020 ). Moreover, the predation rate and efficiency also increase with age (Uiterwaal and DeLong 2018, Ranjbar et al., 2020 ). Therefore, we used adult ladybeetles to assess the predation efficiency and functional response of each species. It is notable that we carried out experiments on the leaves of two host plants: pistachio leaves for the common pistachio psyllid and cucumber leaves for the cotton aphid. These plants differ in terms of classification and morphological characteristics, so our results may be influenced by the specific relationship between the prey species and host plant (Eigenbrode et al., 1998 ; Jalali and Ziaaddini, 2017 ). Because M. sexmaculatus is a generalist predator, it does not prefer psyllids or aphids in any habitat or host plant. This finding aligns with previous reports indicating similarity in the effect of different food sources on the efficiency of this predator species (Mirhosseini et al., 2015 ). In Iran, aphidophagous species tend to move towards pistachio trees when aphids are scarce on herbaceous vegetation. However, O. conglobata behaves quite differently, selecting trees infested by the common pistachio psyllid rather than herbaceous vegetation infested with aphids in the vicinity of pistachio orchards (Jalali, 2001 , 2018). Conclusion Our findings suggest that both ladybeetle species are suitable candidates for the biological control of the common pistachio psyllid. However, given that O. conglobata has adapted to pistachio psyllids as a monotypic host for reproduction and growth, it may be the preferable natural enemy for the biological control of this pest. Nevertheless, further research is needed in the laboratory and under field conditions to determine the effects of biological and environmental factors such as temperature, spatial scale, interactions between the two ladybeetle species (and with other predators), and the impact of chemical pesticides on their performance. Declarations Ethics approval and consent to participate Not applicable Consent for publication Not applicable Availability of data and materials All data are presented in the manuscript. Competing interests The authors declare no competing interests. Funding This work was funded by Rafsanjan University of Medical Sciences (RUMS), Project No. 400283. Author contributions FR: Conceptualization, data collection, writing original draft. MAJ: Supervision, conceptualization, experimental design, methodology. ZA: Visualization, data analysis. XP: Results interpretation, writing and editing. ALM: Conceptualization, writing and editing. TU: writing and editing. 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Biocontrol Sci Technol 14(7):701–711. https://doi.org/10.1080/09583150410001682377 Mehrnejad MR, Jalali MA, Mirzaei R (2011) Abundance and biological parameters of psyllophagous coccinellids in pistachio orchards. J Appl Entomol 135(9):673–681. https://doi.org/10.1111/j.1439-0418.2010.01577.x Mehrnejad MR, Vahabzadeh N, Hodgson CJ (2015) Relative suitability of the common pistachio psyllid, Agonoscena pistaciae (Hemiptera: Aphalaridae), as prey for the two-spotted ladybird, Adalia bipunctata (Coleoptera: Coccinellidae). Biol Control: 80: 128–132. https://doi.org/10.1016/j.biocontrol.2014.10.005 Meseguer R, Levi-Mourao A, Pons X (2021) Species complex and temporal associations between coccinellids and aphids in alfalfa stands in Spain. Insects: 12(11): 971. https://doi.org/10.3390/insects12110971 Mirhosseini MA, Hosseini MR, Jalali MA (2015) Effects of diet on development and reproductive fitness of two predatory coccinellids (Coleoptera: Coccinellidae). Eur J Entomol 112(3):446. https://doi.org/10.14411/eje.2015.051 ISSN 1210 – 575 Obrycki JJ, Kring TJ (1998) Predaceous Coccinellidae in biological control. Annu Rev Entomol 43(1):295–321. https://doi.org/10.1146/annurev.ento.43.1.295 Omkar PA (2005) Ecology of two-spotted ladybird, Adalia bipunctata : a review. J Appl Entomol 129(9–10):465–474. https://doi.org/10.1111/j.1439-0418.2005.00998.x Pervez A, Kumar R, Chandra S (2022) Cumulative functional responses of larvae and adults of two aphidophagous ladybirds. Int J Trop Insect Sci 42(2):1569–1577. https://doi.org/10.1007/s42690-021-00678-2 Rana JS (2006) Response of Coccinella septempunctata and Menochilus sexmaculatus (Coleoptera: Coccinellidae) to their aphid prey, Lipaphis erysimi (Hemiptera: Aphididae) in rapeseed-mustard. Eur J Entomol 103(1):81. https://doi.org/10.14411/eje.2006.013 Ranjbar F, Michaud JP, Jalali MA, Ziaaddini M (2020) Intraguild predation between two lady beetle predators is more sensitive to density than species of extraguild prey. Biocontrol 65:713–725. https://doi.org/10.1007/s10526-020-10036-9 Rogers D (1972) Random search and insect population models. J Anim Ecol 41(2):369–383. https://doi.org/10.2307/3474 Sabaghi R, Hosseini R, Sahragard A (2011) Functional and numerical responses of Scymnus syriacus Marseul (Coleoptera: Coccinellidae) to the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae) under laboratory conditions. J Plant Prot Res 51(4):423–428. https://doi.org/10.2478/v10045-011-0070-4 Saleem M, Hussain D, Anwar H, Saleem M, Ghouse G, Abbas M (2014) Predation Efficacy of Menochilus sexmaculatus Fabricus (Coleoptera: Coccinellidae) against Macrosiphum rosae under laboratory conditions. J Entomol Zool Stud 2(3):160–163 Sanati S, Goldasteh S, Shirvani A, Rashki M (2020) Functional response of Oenopia conglobata contaminata (Coleoptera: Coccinellidae) to Agonoscena pistaciae (Hemiptera: Psyllidae) at two different temperatures. Int J Trop Insect Sci 40:621–628. https://doi.org/10.1007/s42690-020-00108-9 Solomon ME (1949) The natural control of animal populations. J Anim Ecol 1–35. https://doi.org/10.2307/1578 Sugiura K (1998) Suitability of seven aphid species as prey of Cheilomenes sexmaculata (Fabricius)(Coleoptera: Coccinellidae). Jpn J Appl Entomol Zool 42:7–14 Tedders WL, Schaefer PW (1994) Release and establishment of Harmonia axyridis (Coleoptera: Coccinellidae) in the southeastern United States. Entomol News 105(4):228–243 UiterwaalSF, DeLong JP (2018) Multiple factors, including arena size, shape the functional responses of ladybird beetles. J Appl Ecol 55(5):2429–2438. https://doi.org/10.1111/1365-2664.13159 Yuliadhi KA, Supartha IW, Wijaya IN, Pudjianto, Nurmansyah A, Susila IW, Yudha IKW, Utama IWEK, Wiradana PA (2021) The preference and functional response of Sycanus aurantiacus (Hemiptera: Heteroptera: Reduviidae) on three prey types in laboratory conditions. Biodiversitas J Biol Divers 22(12):5662–5667. https://doi.org/10.13057/biodiv/d221252 Cite Share Download PDF Status: Published Journal Publication published 23 May, 2024 Read the published version in Journal of Plant Diseases and Protection → Version 1 posted Reviewers invited by journal 17 Jan, 2024 Editor invited by journal 17 Jan, 2024 Editor assigned by journal 17 Jan, 2024 First submitted to journal 16 Jan, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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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-3871729","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":267739160,"identity":"7362a818-5566-4231-8f5e-236aa749dca7","order_by":0,"name":"Fateme Ranjbar","email":"","orcid":"","institution":"Vali-e-Asr University of Rafsanjan","correspondingAuthor":false,"prefix":"","firstName":"Fateme","middleName":"","lastName":"Ranjbar","suffix":""},{"id":267739161,"identity":"3a1ad88d-1635-48ea-a389-9d24f6f50e48","order_by":1,"name":"M. Amin Jalali","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIiWNgGAWjYBACxgbmBoYHBhJ29v3NB4A8orQAlSUYWCQbSBxLIE4LSBNDAkMF4waGHAPitDC3H2z8kFAgwWzOcOabxM8dNnIM7IePbsBrR09is0SCgQSfZXPvNsneM2nGDDxpaTfwOyuxAaSFmeHA2W0SvG2HgVweM/xa+h82/wBqYWw4kPNM8i9RWmYktoFsYdxwIIdNmjhbZjxsswBqSZaccczYWrYtzZiNkF8M+5MP3/jwp86On7/54c23bTZy/OyHj+HX0oBgs0iASDZ8ykFAHonN/IGQ6lEwCkbBKBiZAAAXwU5ovSLVkgAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0003-4034-541X","institution":"Vali-e-Asr University of Rafsanjan","correspondingAuthor":true,"prefix":"","firstName":"M.","middleName":"Amin","lastName":"Jalali","suffix":""},{"id":267739162,"identity":"569bd08c-0581-4ff6-8605-02ba820cb05e","order_by":2,"name":"Zahra Ahmadi","email":"","orcid":"","institution":"Rafsanjan University of Medical Sciences","correspondingAuthor":false,"prefix":"","firstName":"Zahra","middleName":"","lastName":"Ahmadi","suffix":""},{"id":267739163,"identity":"8b817679-d81c-43d7-b907-a39ba2d08805","order_by":3,"name":"Xavier Pons","email":"","orcid":"","institution":"University of Lleida: Universitat de Lleida","correspondingAuthor":false,"prefix":"","firstName":"Xavier","middleName":"","lastName":"Pons","suffix":""},{"id":267739164,"identity":"ec3be631-fdf9-41d0-826e-8d8dcc69ce21","order_by":4,"name":"Alexandre Levi-Mourao","email":"","orcid":"","institution":"University of Lleida: Universitat de Lleida","correspondingAuthor":false,"prefix":"","firstName":"Alexandre","middleName":"","lastName":"Levi-Mourao","suffix":""},{"id":267739165,"identity":"89c1ce3d-59a4-4492-9c54-9cbbe3708d43","order_by":5,"name":"Todd Ugine","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Todd","middleName":"","lastName":"Ugine","suffix":""}],"badges":[],"createdAt":"2024-01-17 03:34:18","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3871729/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3871729/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s41348-024-00936-8","type":"published","date":"2024-05-24T00:36:04+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":49860793,"identity":"8d5341ed-7243-480f-8409-45a92376b69e","added_by":"auto","created_at":"2024-01-19 08:46:15","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":33675,"visible":true,"origin":"","legend":"\u003cp\u003eObserved numbers of third and fourth instar \u003cem\u003eAphis gossypii\u003c/em\u003e and \u003cem\u003eAgonoscena piataciae\u003c/em\u003e nymphs killed in 24 h by adult females of the ladybeetles \u003cem\u003eOenopia conglobata\u003c/em\u003e (top) and \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e (bottom) \u003cem\u003evs\u003c/em\u003e numbers predicted by the random predator equation (dotted lines).\u003c/p\u003e","description":"","filename":"Slide1.png","url":"https://assets-eu.researchsquare.com/files/rs-3871729/v1/c9d34c8147e23d195a13f92a.png"},{"id":49860109,"identity":"7e97eec0-8980-4585-9446-effa5970f51c","added_by":"auto","created_at":"2024-01-19 08:38:15","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":6575,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between the attack rates of two predatory ladybeetles (\u003cem\u003eOenopia conglobata \u003c/em\u003eand \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e)\u003cem\u003e \u003c/em\u003efeeding on two natural prey (\u003cem\u003eAphis gossypii \u003c/em\u003eor\u003cem\u003eAgonoscena piataciae\u003c/em\u003e). Data are means ± standard deviations. Overlapping error bars have been omitted for clarity.\u003c/p\u003e","description":"","filename":"Slide2.png","url":"https://assets-eu.researchsquare.com/files/rs-3871729/v1/a55f84e9399bf70f9ba80bb6.png"},{"id":57115347,"identity":"04a51c60-b77a-45d4-ae57-6f8a5d3f576a","added_by":"auto","created_at":"2024-05-25 00:36:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":677122,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3871729/v1/eb9835a5-e6e8-46a6-a1f4-d256b705f3e3.pdf"}],"financialInterests":"","formattedTitle":"Comparison of the predatory impacts of indigenous and adventive ladybeetle species (Coleoptera: Coccinellidae) using a functional response approach","fulltext":[{"header":"Introductions","content":"\u003cp\u003ePistachio is one of the most economically important crops in Iran, covering\u0026thinsp;~\u0026thinsp;420,000 ha of land (Mehrnejad, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The common pistachio psyllid \u0026ndash; \u003cem\u003eAgonoscena pistaciae\u003c/em\u003e Burckhardt and Lauterer (Hemiptera: Psyllidae) \u0026ndash; poses a significant threat to cultivated pistachio trees in the Russian Federation, Middle East and Mediterranean areas (Mehrnejad et al. 2010; \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This insect has also evolved resistance to diverse pesticides, making chemical control strategies less effective (Mehrnejad, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Biological control agents offer a promising alternative as part of integrated pest management systems, particularly as a strategy to combat insecticide-resistant pests (Bale et al. 2008). Coccinellids (Col. Coccinellidae), also known as ladybeetles, ladybugs or ladybirds, are beneficial insects in agricultural ecosystems because they help to maintain equilibrium and biological control for a wide range of pests (Obrycki and Kring \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Biddinger et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Kundoo and Khan \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Several predatory ladybeetle species are found in Rafsanjan, the most important pistachio-growing region in Iran (Mehrnejad, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Predatory ladybeetles, along with other biological control agents in the area, control the population of the common pistachio psyllid, which increases rapidly from the emergence of shoots in early spring to early summer and remains stable on pistachio trees until the autumn leaf drop. Nymphs and adult stages of the common pistachio psyllid feed on sap leaves, excreting significant amounts of honeydew that, when dry, lead to leaf and fruit scorching. This feeding behavior has severe economic consequences, such as reduces plant vigor, causing leaf drop, stunting, low fruit yields and the shedding of shoots (Mehrnejad, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). During spring, aphids such as \u003cem\u003eAphis gossypii\u003c/em\u003e Glover (cotton aphid) and \u003cem\u003eAphis craccivora\u003c/em\u003e Koch (cowpea aphid) become established on herbaceous plants earlier than the common pistachio psyllid (Jalali, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Adult ladybeetles gradually emerge from their overwintering sites and disperse on the plants at the edges of pistachio orchards. They engage in mating and egg-laying while also feeding heavily on aphids. At this time, the pistachio tree buds have not yet opened, but the adult psyllids, which typically appear from mid-February onwards, have already infested the pistachio branches. As the common pistachio psyllid begins egg-laying on the swollen buds of pistachio trees, the adult coccinellids begin to feed on these pests.\u003c/p\u003e \u003cp\u003e \u003cem\u003eOenopia conglobata\u003c/em\u003e (Linnaeus) is the most abundant ladybeetle in pistachio orchards. It shows greater activity on pistachio trees even when understory plants such as \u003cem\u003eAlhagi camelorum\u003c/em\u003e are infested with aphids. Moreover, when the pistachio psyllid population is not available in sufficient numbers on pistachio trees, \u003cem\u003eO. conglobata\u003c/em\u003e feeds on aphids infesting understory plants (Jalali, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Mehrnejad and Jalali \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e (Fabricius) is another important predator of aphids, psyllids, thrips, whiteflies, moth larvae, and mites (Sugiura, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Saleem et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Both predators are important biological control agents in pistachio orchards, but \u003cem\u003eO. conglobata\u003c/em\u003e is indigenous whereas \u003cem\u003eM. sexmaculatus\u003c/em\u003e is recently adventive (Jalali, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Ranjbar et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Each species readily feeds on \u003cem\u003eA\u003c/em\u003e. \u003cem\u003epistaciae\u003c/em\u003e and \u003cem\u003eA. gossypii\u003c/em\u003e (Mehrnejad and Jalali, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne of the most important behavioral characteristics of natural enemies from a pest management perspective is the attack rate (\u003cem\u003ea\u003c/em\u003e) against prey (Holling, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1966\u003c/span\u003e). However, this is limited by the time needed by the predator to subdue and consume prey, also known as the handling time (\u003cem\u003eT\u003c/em\u003e\u003csub\u003e\u003cem\u003eh\u003c/em\u003e\u003c/sub\u003e). Both parameters determine the functional response of a natural enemy against prey (Holling, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1966\u003c/span\u003e). This predicts how the predatory behavior of a natural enemy will develop when the prey density increases, and can be used to determine the potential of natural enemies as biological control agents.\u003c/p\u003e \u003cp\u003eThree types of functional responses were first described in the mid-twentieth century (Solomon \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1949\u003c/span\u003e). In a type I response, the predator\u0026rsquo;s consumption rate increases in linearly with the prey density up to the satiation point, whereupon consumption levels plateau and stabilize. In a type II response, the predation rate declines and approaches an asymptote as prey density increases. This occurs because predators are constrained by the time required to handle prey. Finally, a type III response is characterized by an increasing predation rate that correlates positively with prey density. However, there is an inflection point, resulting in an asymptote, when the prey density surpasses the capacity to handle additional prey (Holling \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1959a\u003c/span\u003e). When assessing functional responses, it should be noted that the type of response and its parameters can be influenced by factors such as temperature, host plant, prey life stage, the age, sex and developmental stage of the predator, and the spatial scale (McCaffrey and Horsburgh \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Farhadi et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Jalali et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Madadi et al., 2012; Jalali and Ziaaddini \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Uiterwaal and DeLong 2018; Sanati et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Yuliadhi et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Juliano et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDespite the important role of \u003cem\u003eO. conglobata\u003c/em\u003e and \u003cem\u003eM. sexmaculatus\u003c/em\u003e in pistachio orchards, their functional responses to the common pistachio psyllid population are unknown. We therefore investigated the functional responses of both ladybeetle species against the common pistachio psyllid and the cotton aphid. This will help to determine the value of these ladybeetle species in biological control strategies, taking into account any potential interference caused by the cooccurrence of the two pests. Our data provide valuable insights into the effectiveness and behavior of these ladybeetle species as biological control agents in pistachio orchards, and can be used in the future to optimize pest management strategies.\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eInsect colonies\u003c/h2\u003e \u003cdiv id=\"Sec4\" class=\"Section3\"\u003e \u003ch2\u003ePredatory coccinellids\u003c/h2\u003e \u003cp\u003e \u003cem\u003eM. sexmaculatus\u003c/em\u003e and \u003cem\u003eO. conglobata\u003c/em\u003e colonies were established from several collections of adults made in pistachio orchards of Rafsanjan during the spring season 2022. Individuals were found on pistachio trees infested with the common pistachio psyllid or on understory plants feeding on aphids such as \u003cem\u003eA. gossypii\u003c/em\u003e and \u003cem\u003eA. craccivora\u003c/em\u003e. The orchards selected in collection sites had not been treated with pesticides. The ladybeetles were transported in plastic containers with suitable ventilation to an insect rearing room (26\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 65% \u0026plusmn; 5% relative humidity, 16:8 L:D photoperiod). After sorting and identifying the species, adult specimens were transferred to larger plastic containers (23 \u0026times; 18 \u0026times; 8 cm) with appropriate ventilation. To increase the population, male and female mating pairs were placed in 12-cm Petri dishes lined with paper towels and vented with mesh-covered holes in the lid. To minimize any potential effect of conditioning caused by the feeding on a specific type of prey, each ladybeetle species was reared on a diet of Mediterranean flour moth (\u003cem\u003eEphestia kuehniella\u003c/em\u003e Zeller) eggs for five generations prior to their use in the experiments. For the mating pairs in Petri dishes, we provided two small plastic containers, one filled with moistened cotton wool to provide humidity and the other containing\u0026thinsp;~\u0026thinsp;0.1 g fresh weight of flour moth eggs, which were checked daily (Ranjbar et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). If ladybeetle eggs were found when the Petri dishes were checked, the adults were transferred to new dishes and the containers with the eggs were maintained until the eggs hatched. This management approach allowed for the continuous rearing of ladybeetle populations.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eAphis gossypii\u003c/h2\u003e \u003cp\u003eThe initial population of cotton aphids was collected from infested greenhouses. To establish a colony, squash plants (\u003cem\u003eCucurbita pepo\u003c/em\u003e L.) were used as hosts. Seeds were sown in plastic pots (12 \u0026times; 12.5 cm) containing a 1:1 mixture of cocopeat and perlite before transfer to a growth chamber. The plants were then transferred to mesh cages, which were used to establish a colony of aphids.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eAgonoscena pistaciae\u003c/h2\u003e \u003cp\u003eFresh pistachio leaves infested with common pistachio psyllid were collected from the experimental pistachio garden of Vali-e-Asr University of Rafsanjan, Iran. These insects were used for the functional response experiments within 12 h of collection.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eFunctional response test system\u003c/h2\u003e \u003cp\u003eFunctional responses were tested by exposing mated female \u003cem\u003eM. sexmaculatus\u003c/em\u003e or \u003cem\u003eO. conglobata\u003c/em\u003e (\u0026lt;\u0026thinsp;7 days old, in the process of egg laying) to prey densities of 10, 20, 40, 60, 80, 160 and 200 third and fourth instar nymphs of \u003cem\u003eA. gossypii\u003c/em\u003e or third and fourth instar \u003cem\u003eA. pistaciae\u003c/em\u003e under laboratory conditions (26\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C temperature, 65% \u0026plusmn; 5% relative humidity, 16:8 L:D photoperiod). To standardize the experimental conditions, ladybeetles were starved for 24 h before starting the experiments. We then transferred the nymphs to 12-cm Petri dishes containing a fully mature pistachio leaf (for psyllids) or a 4-day-old cucumber leaf (for aphids). The leaf petiole was placed within an Eppendorf tube filled with water to preserve leaf turgor. Dishes were prepared at different prey densities (For each density, there were 10 biological replicates conducted over the course of a week, with all densities included in one replicate on a single day.) and we counted the number of prey remaining after 24 h.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eFunctional response data analysis\u003c/h2\u003e \u003cp\u003eThe type of functional response was determined by logistic regression, with the ratio of prey consumed as a function of the initial prey density, described by a multinomial function according to Eq.\u0026nbsp;1 (De Clercq et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, Juliano \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2001\u003c/span\u003e):\u003c/p\u003e \u003cp\u003e\u003cimg src=\"data:image/png;base64,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\"\u003e\u003cbr\u003e\u003c/p\u003e \u003cp\u003ewhere \u003cem\u003eN\u003c/em\u003e\u003csub\u003ee\u003c/sub\u003e is the number of prey consumed, \u003cem\u003eN\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e is the initial prey density, and \u003cem\u003eP\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e\u0026ndash;\u003cem\u003eP\u003c/em\u003e\u003csub\u003e3\u003c/sub\u003e are parameters that need to be estimated. The sign (negative or positive) of the linear coefficient in the polynomial function indicates the type of second-order and third-order functional responses, respectively (Juliano, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). The data for the functional response were used as inputs for the random predator equation (Rogers,1972; Juliano, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Equations\u0026nbsp;2 and 3 correspond to the type II and type III functional responses, respectively:\u003c/p\u003e \u003cp\u003e\u003cimg src=\"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAhMAAACwCAIAAAA61nsaAAAAAXNSR0IArs4c6QAAAARnQU1BAACxjwv8YQUAAAAJcEhZcwAADsMAAA7DAcdvqGQAAAASdEVYdFNvZnR3YXJlAEdyZWVuc2hvdF5VCAUAABcUSURBVHhe7d0/bxvJ/cdxd1cliHHPQALUJEUKuzkkQQAfYBVJ48LqXEWFWsOVK0FNcMBBSGnA7gk1aQ56APb1BzdpFLi1U7i7Pv69j9/V/CYz5HKHf1fS+1UInOHu7OzS+/3skpR874skSS1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuTYvWfPnt2b+vrrr3/66aeu966Ko+GhkMbsriTHzz//fHx8/P3333ftMaFWPnr0iBl27WGurq4uLi66xk3GXnz8+LFrTP3www8pOXj88OHDYgFJu3UnkoOi/PTp03HGBpZIDirp6elp17jh2PEXL17k2ZAnByaTyRLJKmlz7kRykBlU564xPkskB7Fxm97MYV+eP3/eNarkAIdotMEv3UG3PzmoyNTlMded1uSgsOZ19nZgj9iveFwnBy+ftx3SeNz+5Pj48ePe3l6qSiPUmhwsP+bdWQ57xH6lx0Vy0MOL6Kcd0ki0JQfn9rxvAaWnsPbLw1W2G8mR33MwwtnZ2cHBwUhuRNiF4UeMyRefGNPDYekOwSyM3y16bZXj2Wrg9FiMVyQmUycHr5TJIY1H8z0HpeT58+ec8HXZjRP+8vKya6/V0tstkoOFqVCYOdRONCUHcy4WpocsiTobO5sW4OeTJ09m7ubSx7PVwOnxmP54XCcHh8jkkMZjmeTgDKfi1Gcyp31cP27C0tvNS1JCbbqhycHCxc6enp6mA1Lv19u3b/MSnGztdRw+vbRrRXLEVIcfIkmb1pwcnM9Pnz6lHFACOMO73uvTO+9Zr1W2Sz26HckROxvlNVCU89/qYI+K41P/tkTYzuvYNL10HFigTo58ryXtVnNycFZzDnO2c61alLBvv/12ZpFai1W2y/JFKaS5YnJQzo6PjxkEDx8+vLq6irlFDxg89UQdZJXz8/ODgwO2/vr1azp56vnz51TwecnBsAzOYizA4xjt73//e/f0/4oKyzL9RyNs/3VcOD2OWDzL3Op3q/JJStqt5uTgBI4qzIP89I5KFI83YentUrAozXkZAmtFce/ajahuh4eH8UlAFPQoeWyLyUShjyRgQzzFMjyOp0DkxHziGvxXv/rVzOSIZ7kt4ClGePHiRYTNvGkzJgv0H42ExZY7nktbOD32K2bCHPIpgZ6Zh0jSTrQlB8UxXZByMudVjIowr6Llohr2mDnIKtulAHGB3zWuFYO0KjbK4zQa1Y0aR5NN8LgIrViSp6KZFv7zn/9clMViSbBRer766qt807nhO7X667iEhdPjqXnJAW7ONjQxSa3akoOTmRM4HkfVi2ttpO/PbMLS22Xhs7OzoihjeJGtxQRYvUDNjQWY1S/3IHt7R0dHbCg6A1tkybwzev70pz/lk2SPKJ3F9Bifzl//+tfzph0LDHkVlj6eq1g4PfYrFpiZHGDOxfGUtBNtycG5nZctHkcd5CTn4rou0OuyynZZJlXJhHUZIR8TMWyPWD6CoVi3EONTkYu5pZl37eslHzx4kC9JkY1S3rWv6/tvf/vb3/zmNzM3HbOqtzgTI+SD8DhmNfx1jFV65ONjyPRYpSc5eLYYU9KuNCQH53zx3kuUA8rc6enp5s7qFbfL6hSsvFgj6vVyc05b79qzMP4333xTb4ImnflkYib5PQd7St0sVozFjo+P66fC8D3ayes4ZHo8Oy85eDww0iRtQUNyzDx7KTdUhPoKcR6qA8v3qIvL6ttlYXSNqeF1thZ1FjzouqadlN14HBP+9OkTiVXMMHafrXft6578c466bvJUDHV5ecl2Z067Hnme1Y/nEoZMj2XiqNZHgOnN3GtJO9GQHJzP9dlLJxVh4Jskyxm4XQoN5WZm7aPuFDOM1ZcuRlFn48u40cNQjMkDCt/R0VGESkwpzxgWKwooQ3311Vf59Fgm3wv6yaSDgwOWiTRilXgqYRn6izBD9KPY9yHHc43mTa+QXiYmUxwB+uu9lrQrQ5Mjvntaf0lp02f1wO3S5DqaWkNtOj4+phn9iCWRd0ahXHrabIVZMUISZTG2ldfliAo6I2OiycLRnEwmNH//+9/n04u5xSAsdnZ29u7dO1ZhthcXF3/961+LfUEM++TJk6IfHJN8Pjt5HXuml2PrMYGZyVHvtSS8f//+5OSka8zy6tWrtf9RqMXJEectZ36oT2DKEKd611ifpu3yIFU9HuTziSqfnqUeUZW6QafywjocJZisihFSVrGV6Ikxo2KGKIXRc35+Hr/fx09eUdbK944Hz6d/UYpVWJJmzJll2Cgj5Auzp7+Mnkl7Glg+jgarDD+e67JweklML44ba+XJAdaKbO7akqa4+nzw4AHhwePPnz8fHR3FicaD6Ix+lulPl1YN71YNRAmIX66uv9G0OVScKDrFY0Ry5D07xDQ4MkWBLpKjH/X04Ppvyi4UrwV5w/jDN5Fs86XkZUpfCK6Tg+NmckgFsmF/f59giCbZ8OOPP/KAn0TF/fv301MRHsRMNFe35uSg1nAlzvw4yYt3jTbql6zoTY61X00vh4mtmBxg+YG7Q/HlVbi4uHj37h11uanybvmlZI/S7UidHPSYHFLh8ePHKQxIi3STAZqUmlevXnXtL18+fPiQZ8mK1pwcnOFcn8Y7OQOr21rkaZE/BiWPupz37AozoTjychaTaU0OSurAmwA2xJKMnNflgbb8UrKttBUe1PccS9wzSbcY2UASdI1ZKDXfffdd15giaYqepa05OaIyxtv32zzP88rIg6LSUXda6+baxa0PByfk4cHcWssidTYvrDMxIMPGoaiPyUKswjy381KyL+k7zaiTg8kUcSvdcScnJyRB16hwb8H5m9+FgNjY39/vGqtZf3LE93Y4z1tL1SoobVwaU2vAg6LSRQ2lNm26Ai5nieQgh168eNG/Cocibk1Y+Ntvv+XuIfoH2uZLyUuTvxNVJAf3495wSAUyoOcGgrOmzhWuAomTDx8+dO0VrDk5ONs555HefduatOlUcXLUnePj43Fet8bVPeZNfiZKbf5fX9RSxecntw5v376N/oG29lKyF3mqxdFIhyImn+eKJHCazEuO+Dy8Toj48IOfXXsFa04OSdIW9CTHyclJ8T5VMDkk6U6blxx0znufwOSQpDttf3//5cuXXeMamZHHBjcf+XtWkRxj/JxDkrQF9XeruNsgGHLFN6lYYKTfrZIkbUHcQHSNL1+4/4i0yBU3JUdHR/VtynJMDkm6kfLfIV8ofod8LW9VweSQpBsp/m7VzK9R1bjhyP8YyYpMDkm6qbjn4E6iPzw+T/+G7snI/1auJGlriI3+Ty+41Vjj3UYwOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVIbk0OS1OZ2JsdkMjk4OHj27FnXltp9/Phxb2/v+Pj46uqq65I0dQuT4/nUp0+fuvauRQG6979++OGH7mmNABcZ3QtzLV12/PTTT4eHh/yMpiTctuTgDCc2usY4kBxct/78889dW6NX/CvyFZQKty05vp/qGuNg3blxiuTgteMV5HXs2tKddwuTY2xvBA1JDpa5uLjoGqP05s2bmbsw/pnX5u1LUt+50vQNKykxOTZuYXLwFLWsa4xYXXBvysxr/eFhckj9TI6NW5gcp6enN6IqMUmm2jWmbsrMa/W+5HjW5JB6mBwb158cdZHKsdajR4/u3bvHz57smSm+L7S3t8cEuq6V5QW0f+bj1xMG9a71LCzdQSbHxvUnBxNG15iDDFi4zEysVf9SCxXw66+/LtKISaavDterJNPJdjPJH6cxa2mZZJUJDDFwMtPpzz6qjGBySD1Mjo3rSQ46nz592l+SoqQusVMMTmmetyIHikqaPztkMjzLMixZLMxobOvq6ipmGxWZZdhxemKZwnITGGLgZNK+dO0MT5kcUg+TY+OoVvOSg6cePnzYX5LYHcrfzPrbX5oZ9uDgYObgrHh2dvbkyZP86r4ul7U0ZjHz9IEzPVzvx0tAz7zPz5eewBADJ8Oz845PPROTQ8qZHBvXkxwUI+pvXfqjn0vy09NTamteXnOM2ZMcHAcuvV+/fh1v3cSld2B8+lmAp1JBHHLo2FbcAM2bOYPMy7nc0hNo0j+ZtC9dO8OUTA6ph8mxcVSoecnBVOvSxi5QTy8vL+Nx8ZZOrj85yBuKe4wzmUzyGs2API7SySbo6R8qiVVYHfXMGYSsmpdzuaUnMNzCyaR96doZk0PqZ3JsHBVqeHLQU5T4vFnoqbZFWSya3MrEWhwuyivj0Oz5lmqSxkExc8SzkQT9lp7AcAsnEwukY5IzOaR+JsfGUaEGJkdd7LhkjsLatafo5Eakli/JyHmTqpfe02crqUbTQzKxMIZU/FRtkc880NmTc8kqE5i37/W9xcLJpH3p2hnWMjmkHibHxlGh5iUHxSj/tICZ5+U4qmpPPWXMefccrJUXU5opSBg2HSJ66GdJSvmQypiqbTHzwDh1ztVWmcBwCyeT9qVrZ5iJySH1uG3J8Ze//GXe93l2hQo1Lzl4ivqbSlJe7Ph5dnY278s/gWVmJgf9jJNqYlEiX79+nY9JP/lUzJCZcCFfhxYrzvxuFWKjrNi1p2aW7yETWNHMyRTSvnTtDJ1Fcvzxj3/85z//2TWkO++2JccIrw17kiMKXKrpVOq457i6unrx4sVkMqF5cXEx738WYvWZycERSDcErMvjlAE0i8mwGFvJQ4L5RM2tDybN9Psc+czBbAmbvAc0i/gZMoHVzZxMIe1L187wlPccUo/FycEJwyUh52F+8RhnO53ov7Jrsvq26jOcksSK/UVko5h/zwU100tFkyWp8syWvWB5+rkoji9HzcQyM5OD/vPz8ziSDJhG4Oj9chCrP0nChvKDRjMOV1338x4exOP8BUL+2nF78e7dO3rihRs4AbAi81/in1bPZAr13iX1vyKTQ8oNvefgHOM8zOsvJ+RafuO3tsq28jOctSgc0wIy3uRgtsy5a4wD9ToOV11b88O7cObsMndO//jHP1iy6XtTMTKRs9Ejk+9LoX6qZ2HpDhqUHJSAzf3Gb2HFbdVnOE2uXkebHCje99850mJmcrAjRQD0z5ynDg8PWYtxYsCB4taEu6V5b9Otrt6XnMkh9RuUHJwz1AhOfkpwOn9SfVmvVbZFOTg6OuJn155ikFWSg03HXUu8wUIApPuYuDZPT4EHLB+rpPnT358cEZY9C2wZOxWxXZTLN2/esC9dY6p/5owDHjSVXUbjCE+m3r592/WuW70vuXrCvKaxL5IwKDk4ZziRUmWkh9N75jvsq1tlW9SaWCXHaEsnBwU0zYFyhqiS9EcwpLnFhAmMb775hlVi+ZQo/ckBnh3VV8Liqj8/mBcXF+xI18j0zJyL+th94rzn05oa2y22vl7z9iWpk4PmvI/TpTtoUHJECeABJ3NUT5rD37mOMlSLC9vCcttiMc521Of20snBiukbSmAE5hzjsJVfYuTRIyIhlZjoTDuVtssIC5NDo1InB+ghAut+6Q5anBx54U7VEJu4JFxuW6zFKT3vPfE0TteeGhJmrNL1ZtJM2Gi6KwqRHPUCPDA5bpaZyRFeT3UN6a5anBycQqnspstq6vu8U2sVS2+Lhae3HOu852AVJjCv4jPs4eEhWVJMeHhyRBRp5/773/92L8m1mclBT+vbbtJttTg5uMLKzyIKJYW46SJ6yAV+GLitGDC/3g/r/ZyDFef9jnEKA2YSH2bQWSRHWt17jhunTg6afs4hJQuSYzu/8RsGbosMiMipT2+WX+93q/JgADNhHGbIPKOTx6QFeBCP0/KsG/00R5UczOfi4qJrjMOb6/+LaSTqf1rx0ncN6c7rS464tEdePcFZVJxX4HaBAh01fQnDt5XOYX7yODqT4pxPwyLqePfEYPkIsbnoiT2lJ55i8E+fPvGT6OI4RE9sblTJwTRG9SWuZFThUSdH3SPdZYvfrRqCk4pTa9O/9Bso2cOTY5sofKRFPaVRJceGPqBaHbNKX47YOZND6ree5IjL8I3+0m9CaTY5lsOR2UK0L2081dnkkPqtITmiYk5/53eDv/SbEBjxTtHMk3mHZ/i85PjXv/71u9/97j//+U/X3h3mVkyPJpEfSbwWHPzizTqQnXsD/mZlPb1dIen/9re/dY0pk0PKreeeIwrQ1k77uMWZubk//OEPO/l/FHqK40iSgzqe/9XIyLmY8BqTI8S/h3zYYusz8exIvr9UJ4f/P4eUW09yjMcIrw1Hkhxk28P//b+YQLPni2cUcWooK3btYVhrub9ZyTLzvga9Zd5zSP1Mjo0bSXJwWPI/phLoXCU5YgFuL/JBGPP1Un+zMu7bFi62BSaH1M/k2DgKIpVo52/CUJGLbzxjleSIQs8B//TpEz/Tm4eMxrDxbHQuTKAQq4whOZg/e9Q1pkwOKWdybBwF8VYmx7OprpE5XfbvY7KYySHdCCbHxlEQb1ZykAfTD85L+delWHHmZxJ5SKTBke5Iepgc0k1hcmzcSJKDw7LGzznS/UTXvsaAaTSeZRlyaOCvH5oc0k1hcmzcSJKDaazxu1XkQZ4c//73v+PBKn8fkxVH8t0q5mBySD1Mjo0bSXIwAWp9ERIcq6XvOViR1Vnm/Pw8Rhj4Nyvj3bD6/StGG8nvczATk0M3yGQyefXqVdeY5eXLl+/fv+8a62BybNxIkgMU67xe559nzHzriZ55yRFPsWK6j0mjFZ+msMX8FSFjWJIH9SvFU/n0doiJmRy6KU5OTo6Ojj5//hxNUuT+/fuciY8fP06dxAadPBXN1ZkcGzee5KgL4vaRDXGDUufEeF47k0M3BbcaxEbX+PLl8vIybj5+/PHH/f19wiP6QXjQs647D5Nj48aTHCg+h9g+bjhmJgdHaTx/K9fk0I3ALQV3Eh8+fOja0xuO7tGXL9999x3Pdo0pns2zZBUmx8aNKjmYxtnZ2Q4nM++e482bN/l7XLtlcuhGIBvyG47CycnJy5cvu8a1e/fucTvSNVZwC5MjCtN4jCo5wEx2+J878erUn3NcXFyMJzZQJAdHjFdwVDOUsL+/T3h0jQx3IcQGunaGe46Z/a1uW3IUV7JjMLbk2Ln4LH1sL1OuSA5fQY0Q8TDzBoIe+kGu1M9yF0J/11jBbUsOTu+nT59OJpPxnOfUnb3rP8CejO3G6I5LXwxL4sYIl5eXh4eHvlWlsYmEmPfWU3zDqvgUBNyjsFbXWMFtSw6QGefn5wcHB2O+qtX4ReRzt7GF/+lSatWfHIgFit/zMDkk6e5amBwwOSRJ/y8+57i8vOzalc+fP7NA/W6Vn3NI0t1VfLfq/fv3RAU9ZAYeP35cfyvX71ZJ0p1GSOS/2RdpQXiABzNvR+7fv99zmzKcySFJNxJRUX97qsdkMnnw4EHXWI3JIUk3VYQBEdK152OZff9ulSQJJycnC8ODwGAZYqZrr8zkkKSbLf2J3HlIl/7v77YyOSRJbUwOSVIbk0OS1MbkkCS1MTkkSW1MDklSG5NDktTG5JAktTE5JEltTA5JUhuTQ5LUxuSQJLUxOSRJbUwOSVKLL1/+D3NjqigK4jNpAAAAAElFTkSuQmCC\"\u003e\u003cbr\u003e\u003c/p\u003e \u003cp\u003ewhere \u003cem\u003eN\u003c/em\u003e\u003csub\u003e\u003cem\u003ee\u003c/em\u003e\u003c/sub\u003e is the number of preys eaten, \u003cem\u003eN\u003c/em\u003e\u003csub\u003e0\u003c/sub\u003e is the initial prey density, \u003cem\u003ea\u003c/em\u003e is the attack constant, \u003cem\u003eT\u003c/em\u003e is the total time available (24 h), \u003cem\u003eT\u003c/em\u003e\u003csub\u003e\u003cem\u003eh\u003c/em\u003e\u003c/sub\u003e is the handling time per prey, and \u003cem\u003eb\u003c/em\u003e, \u003cem\u003ec\u003c/em\u003e and \u003cem\u003ed\u003c/em\u003e are constants. By applying asymptotic 95% confidence intervals (95% CI), we could statistically separate the parameter estimates (Juliano \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Analysis of variance (ANOVA) was applied using SPSS for variance analysis and SAS (SAS Institute 1989) for functional response data analysis. One-way ANOVA was used to compare the functional response parameters.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eA negative correlation coefficient was revealed by linear regression analysis (P1\u0026thinsp;\u0026lt;\u0026thinsp;0) for both ladybeetle species, indicating a type II functional response when feeding on third and fourth instar nymphs of \u003cem\u003eA. gossypii\u003c/em\u003e and \u003cem\u003eA. pistaciae\u003c/em\u003e at different densities (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The logistic regression of prey consumed by female ladybeetles at different initial prey densities indicated the significance of the linear portion of the regression equation (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The decline in the number of prey consumed by ladybeetles with increasing prey density revealed the non-linearity of the regression equation (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The attack rate of \u003cem\u003eO. conglobata\u003c/em\u003e was significantly higher against nymphs of \u003cem\u003eA. pistaciae\u003c/em\u003e compared to \u003cem\u003eA. gossypii\u003c/em\u003e. However, the attack rate of \u003cem\u003eM. sexmaculatus\u003c/em\u003e did not differ significantly between the prey species (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The prey handling time for both ladybeetle species was higher when feeding on \u003cem\u003eA. gossypii\u003c/em\u003e compared to \u003cem\u003eA. pistaciae\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In a 1-h time interval, both ladybeetle species captured a greater number (\u003cem\u003eT\u003c/em\u003e/\u003cem\u003eT\u003c/em\u003e\u003csub\u003eh\u003c/sub\u003e) of \u003cem\u003eA. pistaceae\u003c/em\u003e compared to \u003cem\u003eA. gossypii\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eResults of logistic regression analysis of the proportion of \u003cem\u003eAgonoscena pistaciae\u003c/em\u003e or \u003cem\u003eAphis gossypii\u003c/em\u003e nymphs (3rd and 4th instars) eaten by adult females of two predatory ladybeetles, \u003cem\u003eOenopia conglobata\u003c/em\u003e and \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e, on initial prey density.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\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=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePredator\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePrey\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCoefficient\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEstimate\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eChi Squqre\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\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\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. pistaciae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eConstant (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.4147\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.4791\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e84.8955\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;.0001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLinear (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.0418\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.0133\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e9.8208\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0017\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eO. conglobata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuadratic (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.000014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.000109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.0173\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.8955\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCubic (\u003cem\u003ePc\u003c/em\u003e\u003csub\u003e\u003cem\u003e3\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.631E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.768E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1.7204\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.1896\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. gossypii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eConstant (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.2662\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.2600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e75.9737\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;.0001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLinear (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.0139\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.00799\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3.0111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0827\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuadratic (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.00009\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.000071\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1.6465\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.1994\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCubic (\u003cem\u003ePc\u003c/em\u003e\u003csub\u003e\u003cem\u003e3\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.321E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.944E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4.9411\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0262\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. pistaciae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eConstant (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.7194\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.6261\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e83.4486\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;.0001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLinear (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.0748\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.0169\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e19.6006\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;.0001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. sexmaculatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuadratic (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.000297\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.000135\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4.8586\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0275\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCubic (\u003cem\u003ePc\u003c/em\u003e\u003csub\u003e\u003cem\u003e3\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-3.79E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.324E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1.3021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.2538\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. gossypii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eConstant (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.5616\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.3033\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e71.3431\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;.0001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLinear (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.00559\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.00915\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.3733\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5412\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuadratic (\u003cem\u003eP\u003c/em\u003e\u003csub\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.00018\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.000080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5.1177\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0237\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCubic (\u003cem\u003ePc\u003c/em\u003e\u003csub\u003e\u003cem\u003e3\u003c/em\u003e\u003c/sub\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.604E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.124E-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e9.6705\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0019\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 \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eParameters estimated by the random predator equation and corresponding asymptotic 95% confidence intervals (CI) indicating functional response of adult females of \u003cem\u003eOenopia conglobata\u003c/em\u003e or \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e to densities of 3rd -4th instars of two natural preys, \u003cem\u003eAgonoscena pistaciae\u003c/em\u003e and \u003cem\u003eAphis gossypii\u003c/em\u003e.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\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=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eAsymptotic 95% CI\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e \u003cp\u003eAsymptotic 95% CI\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePredator\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePrey\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eType\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ea\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLower\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eUpper\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eT\u003csub\u003eh\u003c/sub\u003e\u003csup\u003e\u0026Dagger;\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eLower\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eUpper\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eT/ T\u003csub\u003eh\u003c/sub\u003e\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003er\u003csup\u003e2 \u0026sect;\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eO. conglobata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. pistaciae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.315\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.280\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.350\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.317\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.315\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.319\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.962\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. gossypii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.172\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.159\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.185\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.340\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.333\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.347\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.961\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eM. sexmaculatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. pistaciae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.274\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.238\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.310\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.287\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.283\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.290\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.985\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eA. gossypii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.229\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.207\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.251\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.296\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.291\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.301\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.981\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;21.554\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;123.263\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3,48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3,48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"11\"\u003e\u003csup\u003e\u0026dagger;\u003c/sup\u003eattack rate expressed as prey consumed per hour. \u003csup\u003e\u0026Dagger;\u003c/sup\u003eT\u003csub\u003eh\u003c/sub\u003e handling time expressed as hour(s). \u003csup\u003e*\u003c/sup\u003eT/ T\u003csub\u003eh\u003c/sub\u003e estimated maximum numbers of 3rd and 4th instar nymphs of \u003cem\u003eAgonoscena pistaciae\u003c/em\u003e or \u003cem\u003eAphis gossypii\u003c/em\u003e, preyed by one \u003cem\u003eOenopia conglobata\u003c/em\u003e or \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e per hour. \u003csup\u003e\u0026sect;\u003c/sup\u003eThe coefficient of determination [1- (residual sum of squares/corrected total sum of squares)].\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"11\"\u003eMeans (\u0026plusmn;\u0026thinsp;SE) within the same column followed by the same letter are statistically equivalent (Tukey\u0026ndash;Kramer HSD test, α\u0026thinsp;=\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWe used logistic regression analysis to determine the type of functional response exhibited by females of the ladybeetle species \u003cem\u003eO. conglobata\u003c/em\u003e and \u003cem\u003eM\u003c/em\u003e. \u003cem\u003esexmaculatus\u003c/em\u003e when feeding on the third and fourth instar nymphs of \u003cem\u003eA. gossypii\u003c/em\u003e and \u003cem\u003eA\u003c/em\u003e. \u003cem\u003epistaciae\u003c/em\u003e. Our calculations revealed a type II response for both predators against both prey types, indicating that the more the prey density increases, the lower the proportion of prey captured. In other words, the predator\u0026rsquo;s response is density dependent and inversely correlated with the number of prey. At low prey densities, the predator spends more time searching for prey, whereas at high prey densities it invests more time in capturing the prey (Holling, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1959b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn many previous studies, a type II functional response has been documented for various species of predatory ladybeetles when interacting with different prey under diverse laboratory conditions (Hasani et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Khan and Mir, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Jafari and Goldasteh \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sabaghi et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Madadi et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Yuliadhi et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Pervez et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In some cases, the abundance of various life stages of predatory ladybeetles was positively correlated with prey densities (Hodek, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Agarwala \u0026amp; Bardhanroy, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Dixon \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Koch, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Rana, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Meseguer et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Therefore, some researchers have questioned the efficiency of predatory insects with type II functional responses as biological control agents.\u003c/p\u003e \u003cp\u003eAlthough the shape of the functional response curve is important, it may not be sufficient as a sole criterion for predicting the success or failure of a predator in a biological control program. Other factors are also relevant, such as the numerical response, the intrinsic rate of population increase, and the competitive ability of the predator (Pervez and Omkar, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). However, some predators with a type II functional response in laboratory experiments have nevertheless achieved success in mass release programs against pests (Foglar et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Tedders and Schaefer, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1994\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn assessing the potential effectiveness of natural enemies, parameters such as the handling time (\u003cem\u003eT\u003c/em\u003e\u003csub\u003e\u003cem\u003eh\u003c/em\u003e\u003c/sub\u003e) and attack rate (\u003cem\u003ea\u003c/em\u003e) should be taken into account. We found that the attack rate of \u003cem\u003eO. conglobata\u003c/em\u003e against the common pistachio psyllid was significantly higher than against the cotton aphid. In contrast, \u003cem\u003eM. sexmaculatus\u003c/em\u003e attacked each prey species at a similar rate. The longest handling times were recorded for \u003cem\u003eO. conglobata\u003c/em\u003e feeding on cotton aphids, followed by the same predator feeding on the common pistachio psyllid, and then \u003cem\u003eM. sexmaculatus\u003c/em\u003e feeding on the cotton aphid, and finally \u003cem\u003eM. sexmaculatus\u003c/em\u003e feeding on the common pistachio psyllid. The two predator species, \u003cem\u003eM. sexmaculatus\u003c/em\u003e and \u003cem\u003eO. conglobata\u003c/em\u003e, exhibited a higher consumption rate of \u003cem\u003eA. gossypii\u003c/em\u003e compared to \u003cem\u003eA. pistaciae\u003c/em\u003e. Moreover, \u003cem\u003eM\u003c/em\u003e. \u003cem\u003esexmaculatus\u003c/em\u003e demonstrated greater predation on both prey species in comparison to \u003cem\u003eO\u003c/em\u003e. \u003cem\u003econglobata\u003c/em\u003e. The mean fresh weight of a mixture containing 500 third and fourth instar nymphs of \u003cem\u003eA. gossypii\u003c/em\u003e and \u003cem\u003eA. pistaciae\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;ten replicates of 50 nymphs each) was estimated to be 0.283\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009 mg and 0.261\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010 mg, respectively (Jalali et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Despite the equivalent biomass of third and fourth instars in both prey species, this result is likely influenced by the larger size of \u003cem\u003eM. sexmaculatus\u003c/em\u003e, which may be associated with higher voracity or variations in nutritional indices of the prey. It appears that aphid individuals contain more energy than psyllids (Ranjbar et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).An increase in handling time reduced the number of aphids or psyllids consumed by a single predator individual in 1 h. The handling time for both predators feeding on the cotton aphid was significantly longer compared to their feeding on the common pistachio psyllid. Therefore, they consumed more psyllids than aphids.\u003c/p\u003e \u003cp\u003eThe fourth instar larvae and adult ladybeetles are considered the key developmental stages for biological control due to their high feeding capacity (Hodek and Honek, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). Given that the release of fourth-instar larvae is challenging under natural conditions due to their vulnerability, adult ladybeetles are more suitable for release. The predation rate of female insects is generally higher than that of males (Ghosh and Agarwala \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Islam et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, the predation rate and efficiency also increase with age (Uiterwaal and DeLong 2018, Ranjbar et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Therefore, we used adult ladybeetles to assess the predation efficiency and functional response of each species. It is notable that we carried out experiments on the leaves of two host plants: pistachio leaves for the common pistachio psyllid and cucumber leaves for the cotton aphid. These plants differ in terms of classification and morphological characteristics, so our results may be influenced by the specific relationship between the prey species and host plant (Eigenbrode et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Jalali and Ziaaddini, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBecause \u003cem\u003eM. sexmaculatus\u003c/em\u003e is a generalist predator, it does not prefer psyllids or aphids in any habitat or host plant. This finding aligns with previous reports indicating similarity in the effect of different food sources on the efficiency of this predator species (Mirhosseini et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). In Iran, aphidophagous species tend to move towards pistachio trees when aphids are scarce on herbaceous vegetation. However, \u003cem\u003eO. conglobata\u003c/em\u003e behaves quite differently, selecting trees infested by the common pistachio psyllid rather than herbaceous vegetation infested with aphids in the vicinity of pistachio orchards (Jalali, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, 2018).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur findings suggest that both ladybeetle species are suitable candidates for the biological control of the common pistachio psyllid. However, given that \u003cem\u003eO. conglobata\u003c/em\u003e has adapted to pistachio psyllids as a monotypic host for reproduction and growth, it may be the preferable natural enemy for the biological control of this pest. Nevertheless, further research is needed in the laboratory and under field conditions to determine the effects of biological and environmental factors such as temperature, spatial scale, interactions between the two ladybeetle species (and with other predators), and the impact of chemical pesticides on their performance.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAll data are presented in the manuscript.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was funded by Rafsanjan University of Medical Sciences (RUMS), Project No. 400283.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFR: Conceptualization, data collection, writing original draft. MAJ: Supervision, conceptualization, experimental design, methodology. ZA: Visualization, data analysis. XP: Results interpretation, writing and editing. ALM: Conceptualization, writing and editing. TU: writing and editing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful to the Department of Plant Protection, Vali-e-Asr University of Rafsanjan, Rafsanjan, Iran, for the use of greenhouse and laboratory facilities. We thank Dr Richard M. Twyman for English editing and improvement of original draft.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAgarwala BK, Bardhanroy P (1997) Oviposition behavior and reproduction efficiency in ladybird beetles (Coccinellidae: Coleoptera): a case study of \u003cem\u003eMenochilus sexmaculata\u003c/em\u003e (Fabr). 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Biodiversitas J Biol Divers 22(12):5662\u0026ndash;5667. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.13057/biodiv/d221252\u003c/span\u003e\u003cspan address=\"10.13057/biodiv/d221252\" 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":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-plant-diseases-and-protection","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jpdp","sideBox":"Learn more about [Journal of Plant Diseases and Protection](https://www.springer.com/journal/41348)","snPcode":"41348","submissionUrl":"https://www.editorialmanager.com/jpdp","title":"Journal of Plant Diseases and Protection","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Agonoscena pistaciae, Aphis gossypii, prey density, Oenopia conglobata, Menochilus sexmaculatus","lastPublishedDoi":"10.21203/rs.3.rs-3871729/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3871729/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eBiological control has long been recognized for its economic and ecological benefits as a pest management approach, but the use of non-native natural enemies has raised concerns about potential risks and unintended consequences. Indigenous predatory ladybeetles, such as \u003cem\u003eOenopia conglobata\u003c/em\u003e, play a crucial role in controlling many sap-sucking pests, including \u003cem\u003eAgonoscena pistaciae\u003c/em\u003e, which is the most destructive pest of pistachio trees in Iran and other pistachio-growing regions worldwide. However, the abundance of \u003cem\u003eOenopia conglobata\u003c/em\u003e has recently decreased due to the presence of an adventive predatory ladybeetle, \u003cem\u003eMenochilus sexmaculatus\u003c/em\u003e. To better understand the potential risks associated with this adventive species, we investigated the functional responses of female adults of \u003cem\u003eM. sexmaculatus\u003c/em\u003e and \u003cem\u003eO. conglobata\u003c/em\u003e against the third and fourth instar nymphs of \u003cem\u003eA. pistaciae\u003c/em\u003e and \u003cem\u003eAphis gossypii\u003c/em\u003e. Our findings revealed that both predators exhibited a type II functional response, with \u003cem\u003eO. conglobata\u003c/em\u003e demonstrating a significantly higher attack rate against \u003cem\u003eA. pistaciae\u003c/em\u003e nymphs than \u003cem\u003eA. gossypii\u003c/em\u003e nymphs. In contrast, prey species did not have a significant effect on the attack rate of \u003cem\u003eM. sexmaculatus\u003c/em\u003e. Notably, \u003cem\u003eM. sexmaculatus\u003c/em\u003e displayed the highest predation rate and voracity against both prey species. These results provide valuable insights into the potential risks of \u003cem\u003eM. sexmaculatus\u003c/em\u003e for indigenous predatory species like \u003cem\u003eO. conglobata\u003c/em\u003e. The fact that \u003cem\u003eM. sexmaculatus\u003c/em\u003e exhibits higher voracity for both prey species than \u003cem\u003eO. conglobata\u003c/em\u003e does suggest that it may pose a threat to the native ladybeetle population, highlighting the need for further research and careful consideration when introducing non-native natural enemies for pest management.\u003c/p\u003e","manuscriptTitle":"Comparison of the predatory impacts of indigenous and adventive ladybeetle species (Coleoptera: Coccinellidae) using a functional response approach","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-01-19 08:38:11","doi":"10.21203/rs.3.rs-3871729/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2024-01-17T20:45:41+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Journal of Plant Diseases and Protection","date":"2024-01-17T09:26:59+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-01-17T07:28:45+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Plant Diseases and Protection","date":"2024-01-16T07:55:21+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-plant-diseases-and-protection","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jpdp","sideBox":"Learn more about [Journal of Plant Diseases and Protection](https://www.springer.com/journal/41348)","snPcode":"41348","submissionUrl":"https://www.editorialmanager.com/jpdp","title":"Journal of Plant Diseases and Protection","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"420f6b7b-a0ac-4a1b-9525-91529cee5d08","owner":[],"postedDate":"January 19th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-05-25T00:36:04+00:00","versionOfRecord":{"articleIdentity":"rs-3871729","link":"https://doi.org/10.1007/s41348-024-00936-8","journal":{"identity":"journal-of-plant-diseases-and-protection","isVorOnly":false,"title":"Journal of Plant Diseases and Protection"},"publishedOn":"2024-05-24 00:36:04","publishedOnDateReadable":"May 24th, 2024"},"versionCreatedAt":"2024-01-19 08:38:11","video":"","vorDoi":"10.1007/s41348-024-00936-8","vorDoiUrl":"https://doi.org/10.1007/s41348-024-00936-8","workflowStages":[]},"version":"v1","identity":"rs-3871729","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3871729","identity":"rs-3871729","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}