Impact of nC24 agricultural mineral oil deposits on the searching efficiency and predation rate of the predatory mite Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae)

Walking activity, walking straightness, walking speed and searching efficiency of the predatory mite Phytoseiulus persimilis Athias-Henriot were measured on French bean leaf discs that were sprayed with either distilled water, or one of 0.25%, 0.50% and 1.00% w/w aqueous emulsions of an n C24 agricultural mineral oil (AMO). There was no significant difference in percentage of time that mites spent walking in the control (water-sprayed) conditions and in any of the oil treatments. Walking paths were significantly straighter in the oil treatments than in the control, but differences among the oil treatments did not differ significantly. Walking speeds in the oil treatments were significantly slower than in the control and decreased with increasing oil concentration. Deposits of oil at all concentrations significantly suppressed searching efficiency in comparison with control, and searching efficiency in the 1.00% oil treatment was significantly lower than in the 0.25% oil treatment. First predation of P. persimilis on AMO-contaminated eggs of two-spotted mite ( Tetranychus urticae Koch) on unsprayed leaf discs was significantly delayed in all oil treatments in comparison with the control. However there was no significant effect on the overall predation rate. In the tests of P. persimilis predation on AMO-contaminated T. urticae eggs on sprayed leaf discs, the number of first predation occurrences in the first hour was significantly lower in 0.50% and 1.00% oil treatments than in the control. Overall predation rates were significantly reduced by oil but they did not differ significantly among the oil treatments.     

A method to assess the effects of pesticides on the predatory mite Phytoseiulus persimilis (Acari Phytoseiidae) in the laboratory

Phytoseiulus persimilis Athias-Henriot (Acari Phytoseiidae) is a major predator of Tetranychus urticae (Acari Tetranychidae). The performance of P. persimilis in controlling T. urticae may be altered by pesticides used to manage other pests. Therefore, knowledge of the side-effects of pesticides is essential for IPM. A number of laboratory methods were suggested to evaluate pesticide side-effects on predatory mites. Most methods assess residual effects only, and a number of them are characterised by high predator escape rates from experimental units. A method aimed at evaluating the topical and residual effects of pesticides on P. persimilis is herein described. Mites were treated by microimmersion and then reared in holding cells, on bean leaves previously dipped in a pesticide solution. Three insecticides (pyrethrins, spinosad and thiamethoxam), an insecticide-acaricide (abamectin), and two fungicides (azoxystrobin and tolylfluanide) were evaluated. The strain of P. persimilis used for evaluation was collected from unsprayed vegetable plants. All the pesticides affected the survival and fecundity of P. persimilis. Pesticides did not affect the egg-hatching of P. persimilis females exposed to pesticides. Pyrethrins and abamectin proved to be more toxic than other pesticides, and thiamethoxam was more toxic than spinosad, azoxystrobin and tolylfluanide. The escape rate from experimental units was lower than 5% in all trials. Additional experiments were performed on P. persimilis eggs by dipping leaves with eggs in the pesticide solution. None of the pesticides affected egg survival. Semi-field trials conducted on potted bean plants obtained results similar to those reported in laboratory trials.     

A novel disease affecting the predatory mite Phytoseiulus persimilis (Acari, Phytoseiidae): evidence for the involvement of bacteria

Adult female Phytoseiulus persimilis Athias-Henriot (Acari, Phytoseiidae) of a laboratory population show drastic changes in foraging behavior, anatomy and life history compared to typical laboratory populations. We demonstrated earlier that the set of characteristic symptoms, called non-responding (NR) syndrome, is transmitted horizontally between and among predator generations via feces and debris deposited by symptomatic females. Here, we prove that bacteria present in feces and debris deposited by symptomatic females are involved in the induction of the NR-syndrome. The potential of predator products to induce the NR-syndrome was assayed by keeping healthy adult female predators during a period of 3 days on prey-infested bean leaves, which had previously been sprayed with an aqueous suspension of feces and debris. The NR-syndrome was clearly induced in those predators that had been exposed to a suspension collected from symptomatic females (incubation time 4-6 days, 93% shrunken females), whereas predators exposed to a suspension collected from non-symptomatic females did not show the NR-syndrome. Moreover, predators from the first group transmitted infectious products themselves already 5 days after the initial exposure, whereas this was not the case for the second predator group. The bioassay used in the present study is important for laboratories and companies as it can be applied for testing the presence of the novel disease in populations of P. persimilis. To investigate the involvement of bacteria in syndrome induction we (1) eliminated bacteria from a feces-and-debris suspension of symptomatic females by passing the suspension through a bacterial microfilter and (2) added the antibiotic tetracycline to a suspension of feces and debris from symptomatic females. A suspension of feces and debris collected from symptomatic females did not induce the NR-symptom after bacteria had been eliminated, whereas an untreated portion of the same suspension did so. Moreover, the NR-syndrome was induced in predators exposed to an aqueous suspension of the residues that had not passed the bacterial filter. A suspension of feces and debris collected from symptomatic females, to which the antibiotic tetracycline had been added, did not induce the NR-syndrome whereas the same suspension did induce all symptoms when no tetracycline was added. These findings prove that bacteria are involved in the induction of the NR-syndrome. The results are discussed in the context of mite pathology and biological control.     

Absence of odour-mediated avoidance of heterospecific competitors by the predatory mite Phytoseiulus persimilis

Arthropods use odours associated with the presence of their food, enemies and competitors when searching for patches. Responses to these odours therefore determine the spatial distribution of animals, and are decisive for the occurrence and strength of interactions among species. Therefore, a logical first step in studying food web interactions is the analysis of behaviour of individuals that are searching for patches of food. We followed this approach when studying interactions in an artificial food web occurring on greenhouse cucumber in the Netherlands. In an earlier paper we found that one of the predators of the food web, the predatory mite Phytoseiulus persimilis Athias-Henriot, used to control spider mites, discriminates between odours from plants with spider mites, Tetranychus urticae Koch, and plants with spider mites plus conspecific predators. The odours used for discrimination are produced by adult prey in response to the presence of predators, and probably serve as an alarm pheromone to warn related spider mites. Other predator species may also trigger production of this alarm pheromone, which P. persimilis could use in turn to avoid plants with heterospecific predators. We therefore studied the response of the latter to odours from plants with spider mites and 3 other predator species, i.e. the generalist predatory bug Orius laevigatus (Fieber), the polyphagous thrips Frankliniella occidentalis and the spider-mite predator Neoseiulus californicus (McGregor). Both olfactometer and greenhouse release experiments yielded no evidence that P. persimilis avoids plants with any of the 3 heterospecific predators. This suggests that these predators do not elicit production of alarm pheromones in spider mites, and we argue that this is caused by a lack of coevolutionary history. The consequences of the lack of avoidance of heterospecific predators for interactions in food webs and biological control are discussed.     

Plant architecture and prey distribution influence foraging behavior of the predatory mite Phytoseiulus persimilis (Acari: Phytoseiidae)

The arrangement, number, and size of plant parts may influence predator foraging behavior, either directly, by altering the rate or pattern of predator movement, or, indirectly, by affecting the distribution and abundance of prey. We report on the effects of both plant architecture and prey distribution on foraging by the predatory mite, Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae), on cucumber (Cucumis sativus L.). Plants differed in leaf number (2- or 6-leafed), and there were associated differences in leaf size, plant height, and relative proportions of plant parts; but all had the same total surface area. The prey, the twospotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae), were distributed either on the basal leaf or on all leaves. The effect of plant architecture on predator foraging behavior varied depending on prey distribution. The dimensions of individual plant parts affected time allocated to moving and feeding, but they did not appear to influence the frequency with which predators moved among different plant parts. Overall, P. persimilis moved less, and fed upon prey longer, on 6-leafed plants with prey on all leaves than on plants representing other treatment combinations. Our findings suggest that both plant architecture and pattern of prey distribution should be considered, along with other factors such as herbivore-induced plant volatiles, in augmentative biological control programs.     

Side-effects of three pesticides on the predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae)

Side-effects of three commonly used pesticides in Iran were evaluated on an introduced strain of the predatory mite. Phytoseiulus persimilis Athias-Henriot, reared for about 10 years without exposure to any pesticides. Application of pesticides was carried out either to detached bean leaves using a Potter tower at 1 mg wet deposit per cm2 or by a hand sprayer on bean plants until run off. According to an EPPO decision making scheme, pirimiphos-methyl was found to be harmful (E=90.8%) and heptenophos harmless (E=-3.7%) to the predatory mite in the residual initial toxicity tests. For determination of the hazard class of malathion a field test was found to be necessary (E=59.8%). Categories of 1, 2, 3 were determined for heptenophos, malathion and primiphos-methyl, respectively, using IOBC classification. Despite being harmful, it is possible to use pirimiphos-methyl 10 days before release of P. persimilis. Investigation of the contribution of both lethal and sub-lethal effects to total impact indicated the dominance of lethality in the case of pirimiphos-methyl, while malathion acted by both mechanisms. Heptenophos did not have negative effects on fecundity of P. persimilis but rather caused a higher rate of fecundity in comparison with the control. The mortality found in the heptenophos test was not significantly different from the control.     

Disease prevalence and transmission of Microsporidium phytoseiuli infecting the predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae)

Isolated colonies of the predatory mite, Phytoseiulus persimilis, were used to gain information regarding prevalence and transmission of Microsporidium phytoseiuli. Two colonies of P. persimilis were reared on spider mite (Tetranychus urticae)-infested bean plants in isolated cages. Disease prevalence of predators from Colony 1 remained relatively low (between 0 and 15%) over 57 weeks of observation whereas disease prevalence of predators from Colony 2 increased over 3 months (from 12 to 100%). Disease prevalence among predators from Colony 1 had increased to 100% 2 months after weekly sampling had ceased for this colony and periodic sampling confirmed that disease prevalence among individuals of both colonies remained at 100%. Microsporidian spores were not detected in randomly chosen samples of T. urticae prey mites that were removed and examined biweekly during this period. Although numerous microsporidian spores were observed in smear preparations of fecal pellets examined by light microscopy, spores were not observed on leaf surfaces or predator feces when examined by SEM. The latter appeared as intact aggregates composed of numerous dumbbell-shaped crystals and it is unlikely that spores are liberated from intact fecal pellets onto leaf surfaces. Vertical transmission of M. phytoseiuli was 100%; horizontal transmission was low (14.3%) and occurred only when immature P. persimilis were permitted to develop in contact with infected immature and adult predators. The mean number of eggs produced per mated pair was highest when uninfected females were mated with uninfected males (63.2 eggs per mated pair). Although mean egg production decreased when one or both parents were infected, not all differences were significant. Male predatory mites did not contribute to infection of their progeny. Results suggest that routine examination of P. persimilis for microsporidian spores is essential for the management of M. phytoseiuli within P. persimilis colonies. Low disease prevalence and lack of obvious disease signs or symptoms, as in the case of M. phytoseiuli, increase the probability that these pathogens will escape notice unless individuals are routinely examined for pathogens.     

Rainforest habitat resistance to the migration of Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae) in south-eastern Queensland

Abstract  This paper tests the hypothesis that habitat differences affect the migratory ability of the Chilean predatory mite, Phytoseiulus persimilis , an introduced biological control agent of the spider mite, Tetranychus urticae . It is suggested that habitat resistance accounts for the species' inability to invade rainforests in south-eastern Queensland, Australia. Like its prey, P. persimilis migrates to distant plants on air currents. To test our hypothesis, populations of the Chilean predatory mite were established on potted bean plants in both remnant rainforest and adjacent open fields, and their migration monitored using sticky traps. Overall it was found that prey populations on leaves were similar in both habitats, but those of predators were about 20% lower in rainforest. However, the numbers of both predators and prey caught on sticky traps in rainforest were about 6% and 25%, respectively, of those caught in open fields, indicating a strongly reduced rate of aerial migration in the forest. The number of P. persimilis caught on the sticky traps increased with increasing populations of predators on foliage. Thus, dense vegetation inhibits the movement of air currents and inhibits colonisation by both predators and, to a lesser extent, spider mites. These results suggest that the inhibition of aerial migration is one reason for lower numbers of P. persimilis in forest habitats, both because its own vagility is restricted, and because its prey is less able to disperse.     

Improved control capacity of the mite predator Phytoseiulus persimilis (Acari: Phytoseiidae) on tomato

The predatory mite Phytoseiulus persimilis is frequently reported to perform poorly on greenhouse tomatoes. As the predators are mass-reared on another host plant (bean), we supposed that they are poorly adapted to tomato, a plant densely packed with poisonous and sticky glandular hairs. This hypothesis was tested by comparing the control capacity of a stain of P. persimilis directly obtained from a mass rearing with the same strain after four generations on tomato. Both strains were released in a tomato crop in two identical compartments of a greenhouse and the population dynamics of prey (a tomato strain of Tetranychus urticae) and predator were recorded at weekly time intervals. It was found that the strain previously exposed to a tomato environment performed better than the unexposed strain: (1) its population increased faster; (2) the prey population declined faster; and (3) the damage to new-grown tomato leaves was considerably lower. To investigate the causes of the difference in performance between the exposed and unexposed strains, oviposition and survival rates were assessed on a diet of two-spotted spider mites on tomato leaf sections. In addition, the unexposed strain was tested on a diet of two-spotted spider mites on bean leaf sections. The difference in oviposition rates of both predator strains was small compare to the overall mean. However, the oviposition rate of the first generation of predators since transfer from bean to tomato dropped to less than half of the original value. Moreover, mortality in the first generation increased from 14% to 89%, whereas it decreased to 0% after four generations. Future research should clarify whether these changes in life history are due to selection or to physiological adaptation.     

A Novel Disease Affecting the Predatory Mite Phytoseiulus persimilis (Acari, Phytoseiidae): 1. Symptoms in Adult Females

Adult female Phytoseiulus persimilis Athias-Henriot (Acari, Phytoseiidae) of one of our laboratory populations showed a lower degree of attraction to herbivore-induced plant volatiles than other laboratory populations. We hypothesized earlier that this consistent change in foraging behavior is a symptom of a disease, as it is a contagious phenomenon. Here we describe more symptoms by comparing mated females of this population (non-responding (NR) population) with mated females of other populations that are strongly attracted to herbivore-induced plant volatiles (responding populations). The most apparent characteristic of the NR population was the presence of numerous dorso-ventrally flattened females (76% of all females). These females had a normal size after mating but shrank during adulthood. Independent of their age, shrunken females did not reproduce and died a few days after shrinking. In addition to these profound differences in short term performance, females from the NR-population showed behavioral changes, including a lower degree of attraction to herbivore-induced plant volatiles, a higher tendency to leave a prey-patch and a lower predation rate. Moreover, about half of the live females of the NR-population carried birefringent dumbbell-shaped crystals in the legs whereas live females of a responding population carried crystals only in the lumen of the Malpighian tubules and the rectum. The symptom ‘crystals in the legs’ was correlated with low reproduction. Energy dispersive X-ray diffraction of these crystals revealed that they contain calcium and phosphorus along with carbon and oxygen. Crystals with comparable elemental compositions and the same characteristic concentric layering are well known in insects, where they are thought to play a major role in detoxification of calcium and heavy metals, and in storage of phosphorus. The fraction of predators carrying a white spot in the distal part of the opisthosoma, due to accumulation of excretory material in the rectum, was the same in both populations. Present results are discussed in the context of mite pathology and biological control.     

Comparison of thermal activity thresholds of the spider mite predators Phytoseiulus macropilis and Phytoseiulus persimilis (Acari: Phytoseiidae)

The lower and upper thermal activity thresholds of the predatory mite Phytoseiulus macropilis Banks (Acari: Phytoseiidae) were compared with those of its prey Tetranychus urticae Koch (Acari: Tetranychidae) and one of the alternative commercially available control agents for T. urticae, Phytoseiulus persimilis Athias-Henriot. Adult female P. macropilis retained ambulatory function (CT_min) and movement of appendages (chill coma) at significantly lower temperatures (8.2 and 0.4 °C, respectively) than that of P. persimilis (11.1 and 3.3 °C) and T. urticae (10.6 and 10.3 °C). As the temperature was raised, P. macropilis ceased walking (CT_max) and entered heat coma (42.7 and 43.6 °C), beyond the upper locomotory limits of P. persimilis (40.0 and 41.1 °C), but before T. urticae (47.3 and 48.7 °C). Walking speeds were investigated and P. persimilis was found to have significantly faster ambulation than P. macropilis and T. urticae across a range of temperatures. The lower thermal activity threshold data indicate that P. macropilis will make an effective biological control agent in temperate climates.     

Effect of prey density on sex ratio of two predacious mites, Phytoseiulus persimilis and Amblyseius womersleyi (Acari: Phytoseiidae)

The sex ratios of two phytoseiid mites, Phytoseiulus persimilis and Amblyseius womersleyi, were observed under various prey conditions. Upon consumption of abundant prey, both phytoseiids produced progeny in a female-biased sex ratio (approximately 0.8 females). When few prey were consumed, the sex ratio was lowered to 0.5 (the unbiased sex ratio). Under the conditions in which the unbiased sex ratio was observed, male and female progeny appeared in an alternating sequence. To determine the change in the sex ratio and the sequence of progeny, the size of eggs deposited by females under various prey conditions was first compared. Survivorship and developmental rate of progeny (immatures) hatched under ample and poor prey conditions were also examined. The eggs deposited under poor prey conditions were smaller than those deposited under ample prey conditions. This is an indication that the phytoseiid females did not invest extra energy into the eggs to secure survival of their progeny under poor prey conditions. The male and female progeny from the small eggs developed slowly, probably due to the small egg size. However, hatchability of the small eggs and survival of the immatures were the same as those of the normal eggs when the immatures were reared under ample prey conditions. The immature survivorship was little affected by the prey consumption rate of their mothers when the immatures were reared under poor prey conditions. We concluded that the sex ratio of phytoseiid mites is not determined by the characteristics of the progeny, but by the nutritious condition of the females.Exp Appl Acarol 22: 709723 © 1998 Kluwer Academic Publishers     

A Novel Disease Affecting the Predatory Mite Phytoseiulus persimilis (Acari, Phytoseiidae): 2. Disease Transmission by Adult Females

Adult female Phytoseiulus persimilis Athias-Henriot (Acari, Phytoseiidae) of one of our laboratory populations (=NR-population), show the following set of symptoms: predators shrink several days after mating, cease egg production and die several days after shrinking, show a lower degree of attraction to herbivore-induced plant volatiles and a shorter choice time in olfactometer tests, have the tendency to leave a prey patch with ample food, may carry excretory crystals in the legs, may cease prey consumption, and have a lower excretion rate. We hypothesized earlier that this characteristic syndrome, called non-responding (=NR-) syndrome, is caused by a pathogen infecting P. persimilis. To further support this hypothesis we here study several transmission modes of the factor causing the NR-syndrome. In all tests we measured size, short-term fecundity, mortality, predator position, response to plant odors and crystal location, thus including 6 of the 9 symptoms known yet. No evidence was found for vertical transmission from parent to offspring. Eggs from symptomatic females of the NR-population mated by males of the NR-population gave rise to normal-sized, well performing predators, when they had been surface sterilized or transferred to a new leaf. However, such eggs gave rise to shrunken females (17%) when left on the leaf where they had been laid. In the latter case transmission via products deposited on the leaf by the mothers was possible. We therefore tested several modes of horizontal transmission by exposing females of a commercial population that never showed the NR-syndrome (=R1-population) to products related to the symptomatic NR-population. No evidence was found for transmission via food or via squashed adult females. However, symptoms were induced in adult females of the R1-population after a 3-day exposure to a live adult female of the NR-population (incubation period=3–7 days, fraction shrunken females=53%) and after a 1-day exposure to feces and debris collected from such females (incubation period=2–4 days, fraction shrunken females=65%). Contact with live females and feces of the R1-population did not induce the syndrome. These results clearly indicate that the NR-syndrome is a contagious phenomenon and that the factor inducing the syndrome is transmitted horizontally among and between generations via feces and debris deposited by symptomatic females. The results are discussed in the context of mite pathology and biological control.     

Jasmonic acid induces the production of gerbera volatiles that attract the biological control agent Phytoseiulus persimilis

Jasmonic acid (JA) is a plant hormone that is involved in the induction of plant defence in response to herbivore attack. We studied the effect of exposure of gerbera leaves to JA on indirect plant defence, i.e. attraction of natural enemies of herbivores. Treatment of gerbera leaves with JA or feeding damage by the herbivorous spider mite Tetranychus urticae, both induced the production of a complex odour blend that attracts the predatory mite Phytoseiulus persimilis. This phytoseiid predator is a very effective biological control agent of the spider mite T. urticae. Comparison of headspace composition of gerbera leaves exposed to either JA or T. urticae revealed a large degree of resemblance, but some quantitative and qualitative differences were recorded. The major chemical group in both treatments is formed by the terpenoids which quantitatively comprised up to 80% of the total odour blend. These terpenoids included (E)-4,8-dimethyl-1,3,7-nonatriene, (E)--ocimene and linalool that are known to attract P. persimilis. Aldehydes, alcohols, esters and ketones, together with nitrogenous compounds formed the remaining constituents of the odour blend. The induction of predator attractants in plants by JA may be applied in biological control programs, which is discussed in this paper.     

Effects of foliar apple trichomes on Galendromus occidentalis (Nesbitt) (Acari: Phytoseiidae): preferences,fecundity and prey consumption

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Potential predation of the exotic Amblyseius swirskii on Euseius concordis (Acari: Phytoseiidae), a predatory mite commonly found in Brazil

Intra/interspecific predation by A. swirskii.

Both Benin and Netherlands populations of Amblyseius swirskii engaged in cannibalism, and in predation on Euseius concordis larvae. Provision of pollen decreased cannibalism considerably, but had a smaller effect on predation on E. concordis. Potential effect of the eventual introduction of A. swirskii for control of Bemisia tabaci in Brazil is discussed.     

联苯肼酯对智利小植绥螨的安全性及二者对二斑叶螨的联合控制作用

【目的】评价联苯肼酯对智利小植绥螨Phytoseiulus persimilis的安全性,以及该药剂与捕食性天敌智利小植绥螨联合使用的效果。【方法】采用室内生物测定与田间试验的方法测定了联苯肼酯对智利小植绥螨成螨和若螨致死率与繁殖能力的影响,以及二者联合使用对茄子上二斑叶螨的控制效果。【结果】结果表明143 mg/L联苯肼酯对智利小植绥螨成螨和若螨的存活和生殖能力均无显著影响,处理后96 h对若螨的最高致死率为2.30%,成螨为2.04%;处理组8 d的平均产卵量为15.08粒/雌,与对照平均产卵量15.45粒/雌无显著差异;处理组所产卵的平均孵化率为98.63%,与对照组平均孵化率98.13%无显著差异;联苯肼酯悬浮剂在143 mg/L浓度下对二斑叶螨的控制效果表现为速效性高于单独使用智利小植绥螨,但持效性低于智利小植绥螨,防效在第22天时开始下降。二者联合使用表现出较好的速效性与持效性,处理后第2天防效达97.35%,第18天时达100%。【结论】联苯肼酯对智利小植绥螨具有极高的安全性,该药剂与智利小植绥螨联合使用对二斑叶螨具有良好的控制效果。     

Prey preference of predatory mite Amblyseius swirskii (Acari: Phytoseiidae) on Tetranychus urticae (Acari: Tetranychidae) and Bemisia tabaci (Hemiptera: Aleyrodidae)

Tetranychus urticae Koch (Acari: Tetranychidae) and Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae) are major pests in greenhouse crops. Recently, Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae) was shown to be an effective biological control agent of both pests. Therefore, the prey preference of A. swirskii was determined using immature stages of T. urticae and B. tabaci in three various treatments based on Manly's β preference index (β). These treatments consisted of immature stages of two prey species (egg, first and second instar nymphs) with densities 12:12, 6:6 and 3:3, respectively, and with 13 replicates. After 24?h starvation, same-aged females of A. swirskii were added to the leaf discs. All experiments were done on bean leaf discs in Petri dishes (8?cm in diameter) in laboratory conditions with 25?±?2°C, 70?±?5% relative humidity and the photoperiod of 16L:8D hours. Comparing the preference indices using t-tests indicates a significant preference of the predator on eggs (t?=?10.80, df?=?24, P?t?=?8.17, df?=?24, P?T. urticae than B. tabaci. Our findings suggest that developmental stages of prey have effect on the prey selection by A. swirskii.