Response of predatory mites with different rearing histories to volatiles of uninfested plants

The behavioural response of the predatory mite Phytoseiulus persimilis to volatiles from several host plants of its prey, spider mites in the genus Tetranychus, was investigated in a Y-tube olfactometer. A positive response to volatiles from tomato leaves and Lima bean leaves was recorded, whereas no response was observed to volatiles from cucumber leaves, or leaves of Solanum luteum and Solanum dulcamara.Different results were obtained for predators that differed in rearing history. Predators that were reared on spider mites (Tetranychus urticae) on Lima bean leaves did respond to volatiles from Lima bean leaves, while predators that had been reared on the same spider mite species but with cucumber as host plant did not respond to Lima bean leaf volatiles. This effect is compared with the effect of rearing history on the response of P. persimilis to volatile allelochemicals of prey-infested plant leaves.     

The influence of host plant species on parasitism of pollen beetles (Meligethes spp.) by Phradis morionellus

We investigated the response of the specialist insect predator Oligota kashmirica benefica (Coleoptera: Staphylinidae) to volatiles from lima bean leaves infested with the spider mite Tetranychus urticae (Acari: Tetranychidae), both in a Y-tube olfactometer and in a field in Kyoto, Japan. Adult male and female predators were significantly more attracted to T. urticae-infested leaves than to clean air. Adult male and female predators were not more attracted to uninfested leaves, artificially damaged leaves, or the spider mites and their visible products when compared to clean air. In a field trap experiment, 12 adult predators were caught in three traps containing T. urticae-infested lima bean plants over 13 days, whereas no adult predators were trapped in three traps containing uninfested lima bean plants during the same period. These results showed that O. kashmirica benefica adults responded to herbivore-induced plant volatiles from T. urticae-infested lima bean leaves under both laboratory and field conditions.     

Interactions among phytophagous mites, and introduced and naturally occurring predatory mites, on strawberry in the UK

In choice test experiments on strawberry leaf disc arenas the phytoseiid mites Neoseiulus californicus and N. cucumeris were more effective than Typhlodromus pyri as predators of the phytophagous mites Tetranychus urticae and Phytonemus pallidus. There were no preferences shown for either prey by any of these predators. In multiple predator leaf disc experiments both Phytoseiulus persimilis and N. cucumeris significantly reduced numbers of T. urticae eggs and active stages; this effect was seen when the two species were present alone or in combination with other predator species. Neoseiulus californicus was less effective at reducing T. urticae numbers, and T. pyri was not effective; no interaction between predator species was detected in these experiments. When T. urticae alone was present as prey on potted plants, P. persimilis and N. californicus were the only phytoseiids to significantly reduce T. urticae numbers. These two predator species provided effective control of T. urticae when P. pallidus was also present; however, none of the predators reduced numbers of P. pallidus. There were no significant negative interactions when different species of predators were present together on these potted plants. In field experiments, releases of both P. persimilis and N. cucumeris significantly reduced T. urticae numbers. However, there was a significant interaction between these predator species, leading to poorer control of T. urticae when both species were released together. These results show the importance of conducting predator/prey feeding tests at different spatial scales.     

Induced Response of Tomato Plants to Injury by Green and Red Strains of Tetranychus Urticae

We studied the induced response of tomato plants to the green strain and the red strain of the spider mite Tetranychus urticae. We focused on the olfactory response of the predatory mite Phytoseiulus persimilis to volatiles from T. urticae-infested tomato leaves in a Y-tube olfactometer. Tomato leaves attracted the predatory mites when slightly infested with the red strain, or moderately or heavily infested with the green strain. In contrast, neither leaves that were slightly infested with green-strain mites, nor leaves that were moderately or heavily infested with the red strain attracted the predators. We discuss the specific defensive responses of tomato plants to each of the two strains.     

Physiological effect of the host plant on the suitability ofTetranychus urticae as prey forPhytoseiulus persimilis [Acari: Tetranychidae,Phytoseiidae]

Weight gain of adult femalesPhytoseiulus persimilis Athias-Henriot was determined after they were fed eggs or adult females ofTetranychus urticae Koch reared on either Lima bean (Phaseolus vulgaris L.) or nightshade (Solanum douglasii Dunal). No significant difference was found when the predator was fed eggs ofT. urticae reared on either host. Predators were heavier when fed adult femaleT. urticae reared on bean. Apparently, a factor determining the prey's suitability is incorporated into its hemolymph or tissues and is not restricted to its gut content.      

Induced production of extrafloral nectar in intact lima bean plants in response to volatiles from spider mite-infested conspecific plants as a possible indirect defense against spider mites

We found that intact lima bean plants increased the secretion of extrafloral nectar (EFN) after exposure to Tetranychus urticae-induced plant volatiles. Predatory mites, Phytoseiulus persimilis, dispersed more slowly from an exposed intact plant than from a control plant (plant exposed to volatiles from intact conspecific). The predators also dispersed more slowly from those plants that were provided with extra EFN than from untreated plants. We further show that EFN was a potential alternative food source for P. persimilis. From these results, we concluded that increased EFN was involved in the slow dispersal of P. persimilis from the plants exposed to herbivore-induced plant volatiles. Our data suggest that the increase of EFN in an HIPV-exposed intact plant could be an induced indirect defense against spider mites.     

Altered host plant preference of Tetranychus urticae and prey preference of its predator Phytoseiulus persimilis (Acari: Tetranychidae, Phytoseiidae) on transgenic Cry3Bb-eggplants

The behavior of the two-spotted spider mite, Tetranychus urticae Koch and the predatory mite Phytoseiulus persimilis A.-H. was investigated in laboratory experiments with transgenic Bt-eggplants, Solanum melongena L., producing the Cry3Bb toxin and corresponding isogenic, non-transformed eggplants. In bitrophic experiments, dual-choice disc tests were conducted to reveal the effects of transgenic eggplants on host plant preference of T. urticae. Adult spider mite females were individually placed on leaf discs (2 cm diameter) and were observed during five days. Females occurred significantly more frequently on transgenic halves on which also significantly more T. urticae eggs were found. The effects of a Cry3Bb-eggplant fed prey on the feeding preference of P. persimilis were investigated in tritrophic experiments. Sixteen spider mite females, eight of which had been taken from transgenic and eight from isogenic eggplants, were offered to well-fed females of P. persimilis and numbers of respective spider mites consumed were registered 12 h later when the predators were offered new spider mites again. This procedure was repeated six times. The results revealed that predatory mites consumed significantly less Bt-fed spider mites than prey that had been raised on control eggplants. These results indicate that eggplants expressing the Cry3Bb toxin for resistance against the Colorado potato beetle are more preferred by spider mites but are less preferred by their predator P. persimilis. Possible consequences of these findings for biological control of spider mites on eggplants are discussed.     

Influence of pesticide treatments on consumption ofTetranychus urticae [Acarina: Tetranychidae] eggs byPhytoseiulus persimilis [Acarina: Phytoseiidae]

Choice and no-choice experiments were conducted on lima bean leaf discs to determine whether pesticide or water treatments would affect consumption ofTetranychus urticae Koch eggs byPhytoseiulus persimilis Athias-Henriot. In choice experiments,P. persimilis consumed significantly more untreated eggs than eggs treated with fenbutatin-oxide (600 ppm), cyhexatin (200 ppm), or water, when tested at 24, 48, or 72 h after treatment. The surfactant Triton X-100 was included in each pesticide suspension and the water-treated control at 0.1%. On the average, treated eggs amounted to only 30% of the total consumed (n=1,440). The water-treatment effect was weaker than the acaracides at 24 h, but effects did not differ at 48 and 72 h. In 24 h no-choice experiments,P. persimilis consumed as many eggs treated with fenbutatin-oxide, cyhexatin, water, or insecticidal soap (7 575 ppm) as untreated eggs. Fluvalinate treatment of prey eggs (89 ppm), however, resulted in a 50% reduction in egg consumption among predators remaining on leaf discs and a 6-fold increase in abandonment of discs as compared with the untreated control. Continuous consumption of fenbutatin-oxide-treated eggs over 5d did not affectP. persimilis oviposition or egg viability.      

Induced defence in detached uninfested plant leaves: effects on behaviour of herbivores and their predators

Summary Induction of plant defence against herbivores may include the attraction by volatile infochemicals of natural enemies of the herbivore. The emitted volatiles that mediate this attraction may also affect the behaviour of the herbivore itself. In this paper we investigate the response of the herbivorous spider miteTetranychus urticae and the predatory mitePhytoseiulus persimilis towards volatiles whose production is induced in detached Lima bean leaves. Detached uninfested Lima bean leaves were incubated on wet cotton wool on which bean leaves infested with spider mites (T. urticae) were present simultaneously or had been present previously. These treatments induce the production of volatile infochemicals in the uninfested bean leaf tissue: predatory mites are attracted and spider mites are deterred. These are the first data on the response of predators and herbivores to plant volatiles whose production was induced in detached uninfested leaves.     

Predation-related odours reduce oviposition in a herbivorous mite

When adult females of the herbivorous mite, Tetranychus urticae, were exposed to the predatory mite, Phytoseiulus persimilis, they laid fewer eggs than females that had not been exposed to P. persimilis when transferred onto a new leaf patch. However, when T. urticae females were exposed to either products of P. persimilis or artificially damaged conspecific eggs on a leaf patch, the number of T. urticae eggs on a new leaf patch did not differ significantly from the control. The reduced oviposition was neither due to the feeding activity on the leaf patch with P. persimilis nor to that on the new leaf patch. There was also no significant difference between the number of T. urticae eggs produced on a new leaf patch following exposure to the odours of a neighbouring leaf patch where there had previously been either P. persimilis or T. urticae adults. However, female T. urticae that had been exposed to odours from neighbouring leaf patches on which both T. urticae and P. persimilis had been placed produced significantly fewer eggs on a new leaf patch than those that had not been exposed to such odours. Neither odours from neighbouring intact leaf patches on which T. urticae eggs were preyed on by P. persimilis, nor odours from a neighbouring Parafilm patch on which T. urticae was preyed on by P. persimilis affected the oviposition of T. urticae. These data suggest that the presence of T. urticae, P. persimilis and a leaf patch are needed for the emission of odours to reduce oviposition in T. urticae.     

Life-history of predatory mites Neoseiulus californicus and Phytoseiulus persimilis (Acari: Phytoseiidae) on four spider mite species as prey, with special reference to Tetranychus evansi (Acari: Tetranychidae)

The commercially available strains of Phytoseiulus persimilis Athias-Henriot, the biological control agent of Tetranychus urticae Koch, perform poorly in the Western Mediterranean, probably because they are not well adapted to local climatic conditions. For that reason, efforts are being focused on the development of a biological control programme using native phytoseiid mites. Four species of red spider mites can be found in vegetable crops in eastern Spain: T. urticae, Tetranychus turkestani Ugarov and Nikolski, Tetranychus ludeni Zacher and the recently introduced Tetranychus evansi Baker and Pritchard. To evaluate their potential role as biological control agents, the present study evaluates the life-history of local populations of Neoseiulus californicus (McGregor) and P. persimilis when fed on T. urticae, T. turkestani, T. evansi, and T. ludeni in the laboratory. Results indicate that N. californicus and P. persimilis are able to feed and complete their development on the four tested red spider mite species. The predators may exhibit a particularly high capacity for population increase when fed on T. urticae, T. turkestani, and T. ludeni, thus may be able to provide effective control of these species in the field. When fed T. evansi, however, predator performance was poor; significant increase in development and preoviposition times, and a reduction in oviposition period and fecundity were recorded. The resultant low capacity for population growth suggests poor ability of the two tested predators to suppress T. evansi populations on commercial crops. It is unlikely therefore that P. persimilis and N. californicus, now being widely used to control T. urticae in greenhouse crops in Central Europe, will be able to halt any spread of T. evansi to greenhouse crops in temperate areas.     

Local and distant prey-related cues influence when an acarine predator leaves a prey patch

Over relatively long distances, the predatory mite Phytoseiulus persimilis is able to detect volatiles produced by bean plants that are infested by its prey, Tetranychus urticae, the twospotted spider mite. Our investigation examined the separate and combined effects of prey, their products, and prey-induced plant volatiles on when P. persimilis left a potential prey host plant. In wind tunnels, we assessed the relative importance of and interaction among local and distant prey-related cues. The examination of local cues included: (1) all local cues (prey eggs, webbing, and prey-induced plant volatiles), (2) food (prey eggs) and webbing only, (3) plant volatiles only, and (4) no prey-related cues. The examination of distant cues involved the presence or absence of prey-induced plant volatiles from upwind plants. External volatile cues, produced by placing prey-infested plants upwind in the wind tunnel, resulted in more predators leaving downwind plants, and leaving sooner, than when clean plants were upwind, regardless of the availability of prey or prey-related cues on the local plant. However, local cues, especially the presence of food/webbing, had a greater effect than distant cues on timing of predator leaving. Predators remained in larger numbers and for longer times on prey-infested plants. However, the presence of either locally-produced plant volatiles or food/webbing alone still reduced the number of predators leaving a plant in the first hour compared to clean plants. After the first hour, the number of predators leaving was primarily driven by the presence of food/webbing. When no food/webbing was available, predators left plants rapidly; if food/webbing was available, some predatory mites remained on plants at least 24 hours. Even if no food/webbing was available, predators presented with local volatiles remained on plants for several hours longer than on clean plants without local volatiles. These small changes in leaving rates may lead to differences in local population dynamics, and possibly regional persistence, of the predator-prey interaction in patchy environments.     

Interactions in a tritrophic acarine predator-prey metapopulation system V: Within-plant dynamics of Phytoseiulus persimilis and Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae)

To investigate the relative contributions of bottom-up (plant condition) and top-down (predatory mites) factors on the dynamics of the two-spotted spider mite (Tetranychus urticae), a series of experiments were conducted in which spider mites and predatory mites were released on bean plants. Plants inoculated with 2, 4, 8, 16, and 32 adult female T. urticae were either left untreated or were inoculated with 3 or 5 adult female predators (Phytoseiulus persimilis) one week after the introduction of spider mites. Plant area, densities of T. urticae and P. persimilis, and plant injury were assessed by weekly sampling. Data were analysed by a combination of statistical methods and a tri-trophic mechanistic simulation model partly parameterised from the current experiments and partly from previous data. The results showed a clear effect of predators on the density of spider mites and on the plant injury they cause. Plant injury increased with the initial number of spider mites and decreased with the initial number of predators. Extinction of T. urticae, followed by extinction of P. persimilis, was the most likely outcome for most initial combinations of prey and predators. Eggs constituted a relatively smaller part of the prey population as plant injury increased and of the predator population as prey density decreased. We did not find statistical evidence of P. persimilis having preference for feeding on T. urticae eggs. The simulation model demonstrated that bottom-up and top-down factors interact synergistically to reduce the density of spider mites. This may have important implications for biological control of spider mites by means of predatory mites.     

Phytoseiulus persimilis response to herbivore-induced plant volatiles as a function of mite-days

The predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae), uses plant volatiles (i.e., airborne chemicals) triggered by feeding of their herbivorous prey, Tetranychus urticae (Acari: Tetranychidae), to help locate prey patches. The olfactory response of P. persimilis to prey-infested plants varies in direct relation to the population growth pattern of T. urticae on the plant; P. persimilis responds to plants until the spider mite population feeding on a plant collapses, after which infested plants do not attract predators. It has been suggested that this represents an early enemy-free period for T. urticae before the next generation of females is produced. We hypothesize that the mechanism behind the diminished response of predators is due to extensive leaf damage caused by T. urticae feeding, which reduces the production of volatiles irrespective of the collapse of T. urticae population on the plant. To test this hypothesis we investigated how the response of P. persimilis to prey-infested plants is affected by: 1) initial density of T. urticae, 2) duration of infestation, and 3) corresponding leaf damage due to T. urticae feeding. Specifically, we assessed the response of P. persimilis to plants infested with two T. urticae densities (20 or 40 per plant) after 2, 4, 6, 8, 10, 12 or 14 days. We also measured leaf damage on these plants. We found that predator response to T. urticae-infested plants can be quantified as a function of mite-days, which is a cumulative measure of the standing adult female mite population sampled and summed over time. That is, response to volatiles increased with increasing numbers of T. urticae per plant or with the length of time plant was infested by T. urticae, at least as long at the leaves were green. Predatory mites were significantly attracted to plants that were infested for 2 days with only 20 spider mites. This suggests that the enemy-free period might only provide a limited window of opportunity for T. urticae because relatively low numbers of T. urticae per plant can attract predators. Leaf damage also increased as a function of mite-days until the entire leaf was blanched. T. urticae populations decreased at this time, but predator response to volatiles dropped before the entire leaf was blanched and before the T. urticae population decreased. This result supports our hypothesis that predator response to plant volatiles is linked to and limited by the degree of leaf damage, and that the quantitative response to T. urticae populations occurs only within a range when plant quality has not been severely compromised.     

Comparison ofTetranychus evansi andT. urticae [Acari: Tetranychidae] as prey for eight species of phytoseiid mites

Eight phytoseiid species were tested to evaluate and compare their potential as predators ofTetranychus evansi Baker & Pritchard andT. urticae (Koch). The study was conducted using arenas of excised nightshade (Solanum douglasii Dunal) and Lima bean (Phaseolus vulgaris L.) leaves infested with eitherTetranychus species. When the prey wasT. evansi, the predatorsAmblyseius californicus (McGregor) andPhytoseiulus persimilis Athias-Henriot from Ventura, California, showed the highest oviposition rates. However, those rates were 4 to 6 times lower than rates for the same predators feeding onT. urticae. OnlyA. californicus and the strain ofP. persimilis from Beni-Mellal, Morocco, had survivorship higher than 50% 8 days after the beginning of the experiment. The results indicated thatT. evansi is an unfavorable prey for all the phytoseiids tested.

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Résumé Huit espèces d'Acariens phytoseiides étaient testées en vue de comparer leurs potentialités prédatrices vis-à-vis deTetranychus evansi Baker & Pritchard et deTetranychus urticae (Koch). L'étude était réalisée sur des feuilles détachées deSolanum douglasii Dunal et dePhaseolus vulgaris L. Les feuilles avaient été infestées avec l'une ou l'autre des espèces deTetranychus. Amblyseius californicus (McGregor) etPhytoseiulus persimilis Athias-Henriot, de Ventura, ont donné les taux d'oviposition les plus élevés avecT. evansi, mais ces taux étaient 4 à 6 fois inférieurs à ceux obtenus lorsque la proie étaitT. urticae. SeulsA. californicus et la souche deP. persimilis de Beni-Mellal avaient une survie supérieure à 50%, 8 jours après le début du test. Les résultats indiquent queT. evansi n'est pas une proie favorable pour toutes les espèces testées.

     

Functional response ofAllothrombium pulvinum deutonymphs (Acari: Trombidiidae) on two-spotted spider mites (Acari: Tetranychidae)

Functional responses of deutonymphs of the predatory mite,Allothrombium pulvinum Ewing, on eggs and adult females of two-spotted spider mite,Tetranychus urticae Koch, were determined in the laboratory. Predation experiments were conducted on lima bean leaf discs over a 24-h period at 25±2°C, 30–50% RH and 24L: 0D photoperiod. Prey densities ranged from 10 to 120T. urticae eggs per disc or 2 to 32 adult females per disc.Allothrombium pulvinum deutonymphs were more effective againstT. urticae eggs than its adult females. The role ofA. pulvinum deutonymphs in integrated and biological pest control is discussed.     

How Predatory Mites Learn to Cope with Variability in Volatile Plant Signals in the Environment of their Herbivorous Prey

When the chemical cues co-occurring with prey vary in time and space, foraging predators profit from an ability to repeatedly associate chemical cues with the presence of their prey. We demonstrate the ability of a predatory arthropod (the plant-inhabiting mite, Phytoseiulus persimilis) to learn the association of a positive stimulus (herbivorous prey, Tetranychus urticae) or a negative stimulus (hunger) with a chemical cue (herbivore-induced plant volatiles or green leaf volatiles). It has been suggested that the rate at which the integration of information becomes manifest as a change in behaviour, differs between categories of natural enemies (parasitoids versus insect predators; specialist versus generalist predators). We argue that these differences do not necessarily reflect differential learning ability, but rather relate to the ecologically relevant time scale at which the biotic environment changes.     

Experience with methyl salicylate affects behavioural responses of a predatory mite to blends of herbivore-induced plant volatiles

Many natural enemies of herbivorous arthropods use herbivore‐induced plant volatiles to locate their prey. These foraging cues consist of mixtures of compounds that show a considerable variation within and among plant–herbivore combinations, a situation that favours a flexible approach in the foraging behaviour of the natural enemies. In this paper, we address the flexibility in behavioural responses of the predatory mite Phytoseiulus persimilis Athias‐Henriot (Acari: Phytoseiidae) to herbivore‐induced plant volatiles. In particular, we investigated the effect of experience with one component of a herbivore‐induced volatile blend: methyl salicylate (MeSA). We compared the responses of three groups of predatory mites: (1) those reared from egg to adult on Tetranychus urticae Koch (Acari: Tetranychidae) on lima bean plants (Phaseolus lunatus L. that produces MeSA), (2) those reared on T. urticae on cucumber (Cucumus sativus L. that does not produce MeSA), and (3) those reared on T. urticae on cucumber in the presence of synthetic MeSA. Exposure to MeSA during the rearing period (groups 1 and 3) resulted in an attraction to the single compound MeSA in a Y‐tube olfactometer. Moreover, exposure to MeSA affected the choice of predatory mites between two volatile blends that were similar, except for the presence of MeSA. Predators reared on lima bean plants preferred the volatile blend from T. urticae‐induced lima bean (including MeSA) to the volatile blend from jasmonic‐acid induced lima bean (lacking MeSA), but predators reared on cucumber preferred the volatile blend from the latter. Predatory mites reared on cucumber in the presence of synthetic MeSA did not discriminate between these two blends. Exposure to MeSA for 3 days in the adult phase, after rearing on cucumber, also resulted in attraction to the single compound MeSA. We conclude that a minor difference in the composition of the volatile blend to which a predatory mite is exposed can explain its preferences between two odour sources.     

Direct and indirect cues of predation risk influence behavior and reproduction of prey: a case for acarine interactions

Little is known about the flow of chemical information fromhigher to lower levels within the animal food chain. However,this information may determine the behavior and distributionof many animals (e.g., that of potential prey) when exposedto direct and indirect cues of predation risk. We used herbivorousspider mites, Tetranychus urticae Koch (Tetranychidae) as amodel to examine the foraging and oviposition decisions thatprey make when exposed to these cues. We conducted laboratorytests to determine if the previous presence of predators (directcues) on leaf discs or the presence of injured conspecifics(indirect cues) alters the distribution of adults and eggs ofT. urticae. When given a choice, after 24 h, fewer adults and eggswere found on leaf discs that had previously contained specialistspider mite predators, Phytoseiulus persimilis Athias-Henriot (Phytoseiidae),than on discs unexposed to predators. Also, more T. urticaeemigrated from predator-exposed discs than from unexposed discs orfrom those that had previously contained nonpredatory mites(Tyrophagus putrescentiae, Acaridae). Finally, fewer T. urticaeforaged and laid eggs on predator-exposed discs or on thosewith artificially damaged conspecifics (eggs or dead adults)than on discs with intact conspecifics. Tetranychus urticaeprobably recognizes infochemicals (kairomones) from its predatorsor cues from injured spider mites and consequently avoids feedingor ovipositing in areas exposed to these cues. Recognition and avoidanceof kairomones from specialist predators by this prey are likelyto be hereditary, but avoidance of injured conspecifics maybe an adaptation to avoid predators that are not inherentlyrecognized. We discuss the behavioral and ecological implicationsof our findings.