Volatile spider-mite pheromone and host-plant kairomone, involved in spaced-out gregariousness in the spider mite Tetranychus urticae

ABSTRACT. From the host plant-spider mite complex Phaseolus lunatus—Tetranychus urticae Koch a volatile chemical is emitted that acts as a kairomone for the predatory mite Phytoseiulus persimilis Athias-Henriot (Sabelis et al. , 1984a). This kairomone is apparently a byproduct of a vital physiological process and/or it has a function in the biology of the spider mite as well.
The spider mite—host plant complex also emits a volatile spider-mite dispersing pheromone. This is shown in the present study where spider mites were introduced into an odour patch on a horizontal screen in a vertical airflow olfactometer. When spider-mite infested leaves of Lima bean are offered, the spider mites walk mainly straight and soon reach the edge of the screen. On the other hand, when clean Lima bean leaves are offered, the mites walk tortuously most of the time and reach the edge of the screen much later. Artificially damaged plants elicit the same response as undamaged plants. Differences in spider-mite behaviour are observed in the vertical airflow olfactometer when odour of either clean or spider-mite infested leaves is offered. A comparison of the behaviour in these two situations with that when no odour was offered suggests that Lima bean leaves emit a volatile kairomone that activates T. urticae and makes them return after losing the stimulus. A Y-tube olfactometer experiment confirms the existence of this kairomone.
At a low ratio of dispersing pheromone to plant kairomone, the spider mites behave as if only kairomone is present, walking mainly tortuously. At a high ratio they disperse. No aggregation-pheromonal effect is observed.
The possibility that the spider-mite dispersing pheromone acts as a kairomone for P. persimilis is discussed.     

Significance of terpenoids in induced indirect plant defence against herbivorous arthropods

Many plants respond to herbivory by arthropods with an induced emission of volatiles such as green leaf volatiles and terpenoids. These herbivore-induced plant volatiles (HIPVs) can attract carnivores, for example, predators and parasitoids. We investigated the significance of terpenoids in attracting herbivores and carnivores in two tritrophic systems where we manipulated the terpenoid emission by treating the plants with fosmidomycin, which inhibits one of the terpenoid biosynthetic pathways and consequently terpenoid emission.
In the 'lima bean' system, volatiles from spider-mite-infested fosmidomycin-treated plants were less attractive to the predatory mite Phytoseiulus persimilis than from infested control plants. In the 'cabbage' system, fosmidomycin treatment did not alter the attractiveness of Brussels sprouts to two Pieris butterflies for oviposition. The parasitoid Cotesia glomerata did not discriminate between the volatiles of fosmidomycin-treated and water-treated caterpillar-infested cabbage. Both P. persimilis and C. glomerata preferred volatiles from infested plants to uninfested ones when both were treated with fosmidomycin.
Chemical analysis showed that terpenoid emission was inhibited more strongly in infested lima bean plants than in Brussels sprouts plants after fosmidomycin treatment.
This study shows an important role of terpenoids in the indirect defence of lima bean, which is discussed relative to the role of other HIPVs.     

Olfactory response of the anthocorid predatory bug Orius sauteri to thrips-infested eggplants

The predator Orius sauteri (Poppius) (Heteroptera: Anthocoridae) is an effective arthropod natural enemy of thrips, especially Thrips palmi Karny (Thysanoptera: Thripidae), a serious pest of vegetables. First, we studied O. sauteri's response to the odour from thrips-infested eggplant [Solamum melongena L. (Solanaceae)] in a laboratory by two-choice experiments using a Y-tube olfactometer. When detached eggplant leaves were used as odour sources, O. sauteri preferred the volatiles from uninfested leaves to clean air. Concerning preferences among differently infested leaves, O. sauteri preferred the volatiles from plants infested with 10–100 thrips per leaf to uninfested leaves, but showed no significant preference for artificially damaged leaves over uninfested leaves. Similar results were obtained when complete plants were tested as the odour source. Second, release and recapture experiments in a greenhouse, a more realistic set of conditions, were conducted to confirm whether a significant preference for infested plants occurred at similar infestation levels as in the laboratory. Trends favouring infested plants were detected at densities of five and 500 thrips per plant; however, at the five thrips per plant, this trend was due to the large deviation seen in infested plants in only one replicate. In light of the low tolerable thrips density of eggplant, it is necessary to confirm whether artificial treatments with chemicals induce the emission of herbivore-induced plant volatiles that can attract O. sauteri and prolong its residence time on the leaf.     

A maternal influence on the conditioning to plant cues of Aphidius colemani Viereck, parasitizing the aphid Myzus persicae Sulzer

Abstract. The offspring of parasitoids, Aphidius colemani Viereck, reared on Brussels sprouts and emerging from Myzus persicae Sulzer on a fully defined artificial diet, show no preferences in a four-way olfactometer, either for the odour of the diet, the odour of Brussels sprouts, or the odour of two other crucifers (cabbage and Chinese cabbage). A similar lack of odour preferences is shown when the host aphids are exposed for parasitization (for 48 h) on cabbage, Chinese cabbage or wheat. However, if parasitization occurs on Brussels sprouts, a weak but statistically highly significant response to Brussels sprout odour is observed. Although as many as 30–35% of the parasitoids show no response to any odour, another 35% respond positively to the odour of Brussels sprout compared with responses to the odours of cabbage, Chinese cabbage or wheat of only approximately 10%. An analagous result is obtained when the parent parasitoids are reared on cabbage. In this case, significant positive responses of their offspring to cabbage odour occur only if the 48-h parasitization has occurred also on cabbage. However, with parasitoids from Brussels sprouts parasitizing the aphids for 48 h also on Brussels sprouts, the offspring subsequently emerging from pupae excised from the mummies show no preference for Brussels sprout odour. Thus, although the Brussels sprout cue had been experienced early in the development of the parasitoids, they only become conditioned to it when emerging from the mummy. Both male and female parasitoids respond very similarly in all experiments. It is proposed that the chemical cue (probably glucosinolates in these experiments) is most likely in the silk surrounding the parasitoid pupa, and that the mother may leave the chemical in or around the egg at oviposition, inducing chemical defences in her offspring to the secondary plant compounds that the offspring are likely to encounter.     

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.     

Do plants use airborne cues to recognize herbivores on their neighbours?

Plants show defensive responses after exposure to volatiles from neighbouring plants infested by herbivores. When a plant’s neighbours host only species of herbivores that do not feed on the plant itself, the plant can conserve energy by maintaining a low defence level. An intriguing question is whether plants respond differently to volatiles from plants infested by herbivores that pose greater or lesser degrees of danger. We examined the secretion of extrafloral nectar (EFN) in lima bean plants exposed to volatiles from cabbage plants infested by common cutworm, two-spotted spider mites, or diamondback moth larvae. Although the first two herbivore species feed on lima bean plants, diamondback moth larvae do not. As a control, lima bean plants were exposed to volatiles from uninfested cabbage plants. Only when exposed to volatiles from cabbage plants infested by spider mites did lima bean plants significantly increase their EFN secretion compared with the control. Increased EFN secretion can function as an indirect defence by supplying the natural enemies of herbivores with an alternative food source. Of the three herbivore species, spider mites were the most likely to move from cabbage plants to lima bean plants and presumably posed the greatest threat. Although chemical analyses showed differences among treatments in volatiles produced by herbivore-infested cabbage plants, which compounds or blends triggered the increased secretion of EFN by lima bean plants remains unclear. Thus, our results show that plants may tune their defence levels according to herbivore risk level.     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

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.     

Attraction of a generalist predator towards herbivore-infested plants

The occurrence and strength of interactions among natural enemies and herbivores depend on their foraging decisions, and several of these decisions are based on odours. To investigate interactions among arthropods in a greenhouse cropping system, we studied the behavioural response of the predatory bug Orius laevigatus (Fieber) (Hemiptera: Anthocoridae) towards cucumber plants infested either with thrips (Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae)) or with spider mites (Tetranychus urticae Koch (Acari: Tetranychidae)). In greenhouse release-recapture experiments, the predatory bug showed a significant preference for both thrips-infested plants and spider mite-infested plants over clean plants. Predatory bugs preferred plants infested with spider mites to plants with thrips. Experience with spider mites on cucumber leaves prior to their release in the greenhouse had no effect on the preference of the predatory bugs. However, this experience did increase the percentage of predators recaptured. Y-tube olfactometer experiments showed that O. laevigatus was more attracted to odours from plants infested with spider mites than to odours from clean plants. Thus, O. laevigatus is able to perceive odours and may use them to find plants with prey in more natural conditions. The consequences of the searching behaviour for pest control are discussed.     

Response of a phytoseiid predator to herbivore-Induced plant volatiles: Selection on attraction and effect on prey exploitation

Bean plants infested with herbivorous spider mites emit volatile chemicals that are attractive toP. persimilis, a predator of spider mites. In Y-tube olfactometer tests we evaluated involvement of a genetic component in predator response to herbivore-induced plant volatiles. Replicated bidirectional selection resulted in a significant increase in attraction after one generation of selection, but no decrease even after three generations of selection, indicating significant, but unbalanced, additive genetic variation in predator perception of, or response to, herbivore-induced plant volatiles. Selected lines responded differently than an unselected population to food deprivation, pointing to an interaction between their internal state and response to plant volatiles. Selected lines also differed from unselected ones in behaviors associated with local prey exploitation, such as residence time, prey consumption, and reproduction. At lower prey densities,P. persimilis from both “+” lines left spider mite-infested leaves more rapidly and consumed fewer prey eggs than an unselected population. Defining olfactory components of predator search behavior is one step in understanding the effect of plant volatiles on predator foraging efficiency. By selecting lines differing in their attraction to herbivore-induced plant volatiles we may experimentally investigate the link between this behavior, predator foraging efficiency, and local and regional predator-prey population dynamics. The impact of significant additive genetic variation in predator response to plant volatiles on evolution in a tritrophic context also remains to be uncovered.     

State-dependent and odour-mediated anemotactic responses of the predatory mite Phytoseiulus persimilis in a wind tunnel

Anemotaxis in the predatory mite Phytoseiulus persimilis (both well-fed and starved), has previously been studied on a wire grid under slight turbulent airflow conditions yielding weak, yet distinct, gradients in wind speed and odour concentration (Sabelis and Van der Weel 1993). Such conditions might have critically influenced the outcome of the study. We repeated these experiments, under laminar airflow conditions on a flat surface in a wind tunnel, thereby avoiding variation in wind speed and odour concentration. Treatments for starved and well-fed mites were (1) still-air without herbivore-induced plant volatiles (HIPV) (well-fed mites only), (2) an HIPV-free air stream, and (3) an air stream with HIPV (originating from Lima bean plants infested by two-spotted spider mites, Tetranychus urticae). Well-fed mites oriented in random directions in still-air without HIPV. In an air stream, starved mites always oriented upwind, whether plant odours were present or not. Well-fed mites oriented downwind in an HIPV-free air stream, but in random directions in an air stream with HIPV. Only under the last treatment our results differed from those of Sabelis and Van der Weel (1993).     

Feeding History Affects the Response of the Predatory Mite Typhlodromus Kerkirae (Acari: Phytoseiidae) to Volatiles of Plants Infested with Spider Mites

The response of adult females of the predatory mite Typhlodromus kerkirae (Acari: Phytoseiidae) to volatiles emitted from bean leaves infested with Tetranychus urticae (Acari: Tetranychidae) or from leaves of Oxalis corniculata infested with Petrobia harti (Acari: Tetranychidae) was studied in the laboratory using a Y-tube olfactometer. Typhlodromus kerkirae females reared from larvae through to adults on T. urticae and pollen of Vicia faba responded to volatiles of bean leaves infested with T. urticae, either when they had a choice between infested and non-infested bean leaves or between bean leaves infested with T. urticae and O. corniculata leaves infested with P. harti. However, they did not respond when they had been reared only on the carotenoid-deficient pollen of V. faba. Female T. kerkirae that had been reared from larva to the tenth day of adult life on T. urticae and subsequently fed for 1 week on V. faba pollen did not respond to volatiles of bean leaves infested with T. urticae. In contrast, those that had been reared on V. faba pollen to the tenth day of adult life and subsequently fed for 1 week on T. urticae responded to volatiles of infested bean leaves.     

Consequences of constitutive and induced variation in plant nutritional quality for immune defence of a herbivore against parasitism

The mechanisms through which trophic interactions between species are indirectly mediated by distant members in a food web have received increasing attention in the field of ecology of multitrophic interactions. Scarcely studied aspects include the effects of varying plant chemistry on herbivore immune defences against parasitoids. We investigated the effects of constitutive and herbivore-induced variation in the nutritional quality of wild and cultivated populations of cabbage (Brassica oleracea) on the ability of small cabbage white Pieris rapae (Lepidoptera, Pieridae) larvae to encapsulate eggs of the parasitoid Cotesia glomerata (Hymenoptera, Braconidae). Average encapsulation rates in caterpillars parasitised as first instars were low and did not differ among plant populations, with caterpillar weight positively correlating with the rates of encapsulation. When caterpillars were parasitised as second instar larvae, encapsulation of eggs increased. Caterpillars were larger on the cultivated Brussels sprouts plants and exhibited higher levels of encapsulation compared with caterpillars on plants of either of the wild cabbage populations. Observed differences in encapsulation rates between plant populations could not be explained exclusively by differences in host growth on the different Brassica populations. Previous herbivore damage resulted in a reduction in the larval weight of subsequent herbivores with a concomitant reduction in encapsulation responses on both Brussels sprouts and wild cabbage plants. To our knowledge this is the first study demonstrating that constitutive and herbivore-induced changes in plant chemistry act in concert, affecting the immune response of herbivores to parasitism. We argue that plant-mediated immune responses of herbivores may be important in the evaluation of fitness costs and benefits of herbivore diet on the third trophic level.     

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.     

Cultivar‐specific plant odour preferences of a generalist aphid parasitoid Aphidius colemani and a possible mechanism for maternal priming of resistance to toxic plant chemistry

Aphidius colemani Viereck, emerging from Myzus persicae (Sulzer) mummies on the Brussels sprout cultivar ‘Bedford Winter Harvest’ (BWH), responds positively in the olfactometer to the odour of that cultivar in comparison with air. Responses to the odours of other sprout cultivars, cabbage and broad bean could be explained by the humidity from plant leaves. In a choice between BWH and other sprout cultivars, the BWH odour is preferred, or that of cv. ‘Red Delicious’ (RD) if the parasitoids are reared on RD. This confirms previous work showing that the secondary chemistry of a cultivar is learnt from the mummy cuticle during emergence. Adults emerging from pupae excised from the mummy show a similar but less pronounced preference. Parasitoids developing in aphids on an artificial diet do not discriminate between the odours of BWH and RD, unless allowed contact with a mummy from the same cultivar that the mother develops on. This suggests a cultivar‐specific maternal cue. This cue is speculated to consist of a small amount of the secondary chemistry (probably glucosinolates in the present study) that are left in or on the egg at oviposition, which subsequently induces enzymes that detoxify plant‐derived toxins in the aphid host. Indeed, when parasitoids emerging from diet‐reared aphids are released on aphid‐infested sprout plants, fewer mummies are produced than by parasitoids emerging from mummies of plant‐reared aphids or from excised pupae. Only parasitoids that emerge from mummies of plant‐reared aphids prefer the cultivar of origin as shown by the number of mummified hosts.     

Odour-mediated preference and prey preference of Macrolophus caliginosus between spider mites and green peach aphids

Abstract  A comparative study between direct prey preference and odour-mediated preference of the predatory mirid bug, Macrolophus caliginosus , was conducted. We used a Y-tube olfactometer to determine the attraction of the predator to herbivore-induced volatiles from Myzus persicae or Tetranychus urticae -infested sweet pepper plants over clean plants and to direct prey odours over clean air. The mirid bugs showed a stronger response to odours from infested plants than to odours from clean plants. The mirids did, however, not seem to exploit odours emitted directly from the prey themselves. Our results further demonstrated that M. caliginosus prefers M. persicae to T. urticae in a direct two-choice consumption test. This preference was, however, not reflected in a similar odour-mediated preference between plant volatiles induced by either of the two preys. Two hypotheses are suggested for this neutral response of the mirids to herbivore-induced volatiles from a preference prey or a non-preference prey.     

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.     

Are herbivore-induced plant volatiles reliable indicators of herbivore identity to foraging carnivorous arthropods?

Plants that are damaged by herbivorous arthropods provide carnivorous enemies of the herbivores with important information. They emit an induced volatile blend that is highly detectable to the carnivores from a distance. Such detectable signals that indicate herbivore presence are important for the carnivores because herbivores themselves are under strong selection not to expose themselves. In addition, carnivores would benefit from a specificity of the induced plant volatiles. Whether herbivore-induced plant volatiles are reliable indicators of herbivore identity, however, has not been resolved unambiguously. Some studies support the reliability of herbivore-induced plant volatiles, while others do not. Different approaches have been used such as chemical analysis, behavioural analysis or a combination of the two. Based on the total of chemical studies one might conclude that in most cases herbivore-induced plant volatiles are not very specific for the herbivore that damages the plant. However, arthropod chemosensors are much more sensitive than the detectors of analytical instruments. Therefore, chemical analyses are not suitable to demonstrate whether or not herbivore-induced plant volatiles are reliable indicators of herbivore identity to carnivores. Behavioural studies should provide this information. In analysing carnivore behaviour it should be realised, however, that arthropod behaviour can be highly variable. Arthropod foraging decisions are affected by external and internal factors such as (a) abiotic environmental factors, (b) presence of competitors or enemies, (c) deprivation of food or oviposition sites, (d) specific deprivation of certain nutrients or (e) learning. In this paper their effect on discrimination of carnivores between volatile blends emitted by plants infested by different herbivores is reviewed. This provides testable hypotheses of why discrimination was not found in some studies. The ability of carnivores to discriminate is likely to be more common than is clear to date, which should invoke functional studies of the conditions that influence the occurrence of this discrimination.     

Conditioned olfactory responses of a predatory mite, Neoseiulus womersleyi, to volatiles from prey-infested plants

To clarify the prey‐finding behavior of the predatory mite Neoseiulus womersleyi (Schicha) (Acari: Phytoseiidae), we studied its olfactory responses to volatiles from the prey‐infested plant on which the mites had been collected. We used a local N. womersleyi population called Kanaya collected from tea (Camellia sinensis L.) (Theaceae) plants infested by Tetranychus kanzawai Kishida (Acari: Tetranychidae) in Kanaya City, Japan. Neoseiulus womersleyi (Kanaya population) were more attracted to volatiles from tea plants infested with five female T. kanzawai per leaf for 7 days than to intact tea leaves in a Y‐tube olfactometer. Tetranychus kanzawai‐induced tea leaf volatiles were identified as (E)‐β‐ocimene, (E)‐4,8‐dimethyl‐1,3,7‐nonatriene, and (E,E)‐α‐farnesene. As olfactory responses are known to differ among local populations of N. womersleyi, we compared the responses of the Kanaya population with those of a Kikugawa population collected from tea plants infested by T. kanzawai in Kikugawa City. To test the influence of previous predation experience, we reared the two populations on tea plants infested by T. kanzawai or on kidney bean plants (Phaseolus vulgaris) infested by Tetranychus urticae Koch. The Kanaya population was more attracted to the volatiles from infested plants on which they had been reared. Because the Kanaya population was not attracted to the plant volatiles they had not previously experienced, the positive response to previously experienced volatiles might be the result of learning. By contrast, the Kikugawa population showed no preference for previously experienced volatiles from infested plants. The implications of this flexibility in foraging behavior are discussed.