Herbivore host plant selection: whitefly learns to avoid host plants that harbour predators of her offspring

Evidence is accumulating that herbivorous arthropods do not simply select host plants based on their quality, but also on the predation risk associated with different host plants. It has been suggested that herbivores exclude plant species with high predation risk from their host range. This assumes a constant, predictable predation risk as well as a rather static behaviour on the part of the herbivore; plants are ignored irrespective of the actual predation risk. We show that adult females of a small herbivore, the whitefly Bemisia tabaci, can learn to avoid plants with predatory mites that attack only juvenile whiteflies, while they accept host plants of the same species without predators. Predatory mites disperse more slowly than whiteflies; they cannot fly and walk from plant to plant. Hence, by avoiding plants with predators, the whiteflies create a temporary refuge for their offspring. We suggest that the experience of arthropod herbivores with risks associated with host plants plays an important role in their host plant selection.     

How predatory mites find plants with whitefly prey

We investigated the searching behaviour of two species of predatory mites, Typhlodromips swirskii (Athias-Henriot) and Euseius scutalis (Athias-Henriot), both known to feed on immature stages of the whitefly Bemisia tabaci Gennadius. When released in a greenhouse inside a circle of cucumber plants that were alternatingly clean or infested with immature whiteflies, the mites took several days to find plants. Both species were recaptured significantly more on plants with whiteflies. This suggests that the mites are able to discriminate between plants with and without whiteflies. The predators may either have been attracted to plants with whiteflies from a distance or arrested on plants with whiteflies. Typhlodromips swirskii that had previously fed on whitefly immatures on cucumber leaves were significantly attracted by volatiles from cucumber plants with whiteflies in a Y-tube olfactometer. This suggests that the mites use volatile cues to discriminate between infested and clean plants. However, this response waned rapidly; if predators, experienced as above, were starved for 3–4 h in absence of cucumber leaves, they no longer preferred volatiles of infested plants to clean plants. Furthermore, T. swirskii that had no experience with immature whiteflies on cucumber plants also did not prefer odours of infested plants to those of clean plants. Because the release experiment with this species in the greenhouse was done with inexperienced predators, this suggests that the aggregation of mites on plants with whiteflies was mainly caused by differential arrestment of mites on plants with prey and clean plants. For T. swirskii, this was in agreement with the finding that the fraction of predators on plants with prey increased with time to levels higher than 70%. A less clear trend was found for E. scutalis, for which the fraction of predators on plants with prey stabilized soon after release to levels from 54–70%. Hence, the predatory mites may find plants with prey by random searching, but they are subsequently arrested on these plants. An earlier study showed that 87% of all whiteflies released in a set-up as used here were recaptured within 1 day. Hence, the effectiveness with which predatory mites locate plants with whiteflies is low compared with that of their prey. We expect this to generate spatial patterns in the dynamics of predator and prey and this may have consequences for biological control of whiteflies with predatory mites.     

Non-consumptive effects of the predatory beetle Delphastus catalinae (Coleoptera: Coccinellidae) on habitat use patterns of adult whitefly Bemisia argentifolii (Hemiptera: Aleyrodidae)

This study examined habitat use patterns by adult whiteflies, Bemisia argentifolii Bellows and Perring, in response to predators, Delphastus catalinae (Horn), at different spatial scales. When female whiteflies were confined to small arenas with leaf discs from which they could not escape, whiteflies significantly delayed settling on leaf discs when predators were present compared to when no predators were introduced. The presence of D. catalinae altered the vertical distribution of adult whiteflies (sex ratio = 1:1) on cucumber plants; adult whiteflies moved upward faster over time within the plant canopy when predators were present mainly on the lower leaves of the plants compared to whiteflies on plants without predators. Most D. catalinae remained in the lower parts of the plants during the experiment. Therefore, we inferred that female whiteflies more quickly moved to the upper plant strata to reduce the risk of predation of their progeny; this would induce subsequent movement of males seeking mates. Introduction of D. catalinae onto a cucumber plant with high whitefly density did not cause increased dispersal of adult whiteflies (sex ratio = 1:1) into neighboring uninfested plants. The results indicate that predator-avoidance behaviors by adult B. argentifolii differed at different spatial scales. The predator-avoidance behavior may have a negative impact at the within-plant scale by inducing more whiteflies to move into upper plant strata. However, the effect of predators on the among-plant dispersal of whiteflies was not significant.     

Pest species diversity enhances control of spider mites and whiteflies by a generalist phytoseiid predator

To test the hypothesis that pest species diversity enhances biological pest control with generalist predators, we studied the dynamics of three major pest species on greenhouse cucumber: Western flower thrips, Frankliniella occidentalis (Pergande), greenhouse whitefly, Trialeurodes vaporariorum (Westwood), and two-spotted spider mites, Tetranychus urticae Koch in combination with the predator species Amblyseius swirskii Athias-Henriot. When spider mites infested plants prior to predator release, predatory mites were not capable of controlling spider mite populations in the absence of other pest species. A laboratory experiment showed that predators were hindered by the webbing of spider mites. In a greenhouse experiment, spider mite leaf damage was lower in the presence of thrips and predators than in the presence of whiteflies and predators, but damage was lowest in the presence of thrips, whiteflies and predators. Whitefly control was also improved in the presence of thrips. The lower levels of spider mite leaf damage probably resulted from (1) a strong numerical response of the predator (up to 50 times higher densities) when a second and third pest species were present in addition to spider mites, and (2) from A. swirskii attacking mobile spider mite stages outside or near the edges of the spider mite webbing. Interactions of spider mites with thrips and whiteflies might also result in suppression of spider mites. However, when predators were released prior to spider mite infestations in the absence of other pest species, but with pollen as food for the predators, we found increased suppression of spider mites with increased numbers of predators released, confirming the role of predators in spider mite control. Thus, our study provides evidence that diversity of pest species can enhance biological control through increased predator densities.     

捕食螨化学生态研究进展

捕食螨是重要的生物防治因子。早在20世纪70年代就发现了捕食螨的性信息素,许多研究证明植物挥发物在捕食螨向猎物定位过程中发挥着至关重要的作用,影响捕食螨寻找猎物的植物挥发物来源于未受害植物、机械损伤植物、猎物危害植物、非猎物危害植物。人工合成的植物挥发物组分对捕食螨具有引诱作用,但引诱活性低于虫害诱导植物释放的挥发性混合物。捕食螨的饲养条件、饥饿程度、学习与经验行为等会影响捕食螨对植物挥发物的反应。介绍了信息素与植物挥发物对捕食螨的作用,并讨论了目前存在的问题和研究前景。     

Previous and Present Diets of Mite Predators Affect Antipredator Behaviour of Whitefly Prey

Predator diet is known to influence antipredator behaviour in prey. Yet, it is not clear how antipredator behaviour is affected by diet changes of the predator. We studied the effect of previous and present diet of a predatory mite Typhlodromips swirskii on the antipredator response of its prey, the whitefly Bemisia tabaci. An earlier study showed that adult female whiteflies that had experienced predators, had learned to avoid ovipositing on plants with predators whose previous and present diet consisted of whitefly eggs and immatures. Here, we investigate whether adult whiteflies also avoid plants with predators whose present and/or previous diet consisted of a non-whitefly food source. Adult whiteflies were found not to avoid plants with predators whose present diet consisted of pollen and whose previous diet had consisted of either pollen or whitefly eggs and larvae. They did avoid plants with predators whose present diet consisted of whiteflies and whose previous diet had consisted of pollen, but to a lesser extent than when previous and present diet consisted of whiteflies. In a choice experiment, whiteflies discriminated between plants with predators whose present diet consisted of whiteflies, but that differed in previous diet. Our results show that both previous and present diets of predators are important in eliciting antipredator behaviour.     

Plants with spider-mite prey attract more predatory mites than clean plants under greenhouse conditions

Although many predators and parasitoids are known to respond to odours produced by plants infested with their prey under laboratory conditions, there are actually few studies that show that this response leads to higher numbers of predators or parasitoids on the plants under natural conditions. Here we study the response of predatory mites (Phytoseiulus persimilis Athias-Henriot, Acari, Phytoseiidae) to odours from cucumber plants infested with two-spotted spider mites (Tetranychus urticae Koch, Acari, Tetranychidae) in greenhouse release experiments, where predators were released in the centre of a hexagon of cucumber plants. Forty to 57% of all predators released were recaptured on plants within 7 h. Of these, an average of 79.5% were found on infested plants, indicating that these attract about 4 times as many predators as do clean plants. Hence, the blind predatory mites were guided to the plants with prey by herbivore-induced odours produced by the plant, as was indicated by olfactometer experiments, where it was found that P. persimilis preferred odours from infested cucumber plants to odours from clean cucumber plants. The long-range searching behaviour of P. persimilis is discussed.     

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.     

Prey preference,intraguild predation and population dynamics of an arthropod food web on plants

The theory of intraguild predation (IGP) largely studies effects on equilibrium densities of predators and prey, while experiments mostly concern transient dynamics. We studied the effects of an intraguild (IG) predator, the bug Orius laevigatus, on the population dynamics of IG-prey, the predatory mite Phytoseiulus persimilis, and a shared prey, the phytophagous two-spotted spider mite Tetranychus urticae, as well as on the performance of cucumber plants in a greenhouse. The interaction of the predatory mite and the spider mite is highly unstable, and ends either by herbivores overexploiting the plant or predators exterminating the herbivores. We studied the effect of IGP on the transient dynamics of this system, and compared the dynamics with that predicted by a simple population-dynamical model with IGP added. Behavioural studies showed that the predatory bug and the predatory mite were both attracted to plants infested by spider mites and that the two predators did not avoid plants occupied by the other predator. Observations on foraging behaviour of the predatory bug showed that it attacks and kills large numbers of predatory mites and spider mites. The model predicts strong effects of predation and prey preference by the predatory bugs on the dynamics of predatory mites and spider mites. However, experiments in which the predatory bug was added to populations of predatory mites and spider mites had little or no effect on numbers of both mite species, and cucumber plant and fruit weight.     

Avoidance of natural enemies by adult whiteflies, Bemisia argentifolii, and effects on host plant choice

In this study we asked whether, in the context of a trap crop system, differential predation risks among plants influence host choice patterns of adult whiteflies, Bemisia argentifolii. We investigated whether adult whiteflies avoid natural enemies inhabiting poinsettia (a cash crop) and whether this behavior can be used to increase the movement of whiteflies to cucumber (a trap crop). The potential of cucumber as a trap crop was first evaluated and we found that significantly more whiteflies were attracted to cucumber when the whiteflies were released between the two plants. However, the accumulation of whiteflies on cucumber substantially diminished if the insects had first settled on poinsettia. Under such circumstances, we investigated whether movement of adult whiteflies to cucumber could be increased by creating conditions that would cause the whiteflies that had settled on poinsettia to leave the plant. A mechanical disturbance, consisting of shaking the plant, was first used to test this hypothesis. The shaking caused more whiteflies to leave poinsettia and move onto the trap crop, compared to undisturbed whiteflies. We then asked whether the presence of natural enemies on the cash crop could induce whiteflies to leave the cash crop and move onto the trap crop. Three natural enemies were tested: two predators, Amblyseius swirskii and Delphastus catalinae, and a parasitoid Encarsia formosa. The presence of D. catalinae on poinsettia induced significantly more whiteflies to disperse to cucumber compared to poinsettia with no natural enemies, whereas A. swirskii and E. formosa did not result in a significant increase. Predator avoidance behavior by adult whiteflies should be investigated further in the context of trap cropping and other crop-habitat alterations designed to help manage whitefly abundance.     

Potential lethal and non-lethal effects of predators on dispersal of spider mites

Predators can affect prey dispersal lethally by direct consumption or non-lethally by making prey hesitate to disperse. These lethal and non-lethal effects are detectable only in systems where prey can disperse between multiple patches. However, most studies have drawn their conclusions concerning the ability of predatory mites to suppress spider mites based on observations of their interactions on a single patch or on heavily infested host plants where spider mites could hardly disperse toward intact patches. In these systems, specialist predatory mites that penetrate protective webs produced by spider mites quickly suppress the spider mites, whereas generalist predators that cannot penetrate the webs were ineffective. By using a connected patch system, we revealed that a generalist ant, Pristomyrmex punctatus Mayr (Hymenoptera: Formicidae), effectively prevented dispersal of spider mites, Tetranychus kanzawai Kishida (Acari: Tetranychidae), by directly consuming dispersing individuals. We also revealed that a generalist predatory mite, Euseius sojaensis Ehara (Acari: Phytoseiidae), prevented between-patch dispersal of T. kanzawai by making them hesitate to disperse. In contrast, a specialist phytoseiid predatory mite, Neoseiulus womersleyi Schicha, allowed spider mites to escape an initial patch, increasing the number of colonized patches within the system. Our results suggest that ants and generalist predatory mites can effectively suppress Tetranychus species under some conditions, and should receive more attention as agents for conservation biological control in agroecosystems.     

Leaf domatia reduce intraguild predation among predatory mites

1. Although theory suggests that intraguild predation destabilises food webs and may result in exclusion of species, empirical observations of food webs reveal that it is a common interaction. It has been proposed that habitat structure reduces the interaction strength of intraguild predation, thus facilitating the coexistence of species. 2. This was tested using acarodomatia, tiny structures on plant leaves, and predatory mites, which usually reside in these domatia. Sweet pepper plants (Capsicum annuum L.) were used, which possess domatia consisting of tufts of hair, and coffee plants (Coffea arabica L.) with pit‐shaped domatia. 3. On sweet pepper, the predatory mites Neoseiulus cucumeris Oudemans and Iphiseius degenerans Berl. feed on each other's juveniles. Larvae of each of the species were therefore used as intraguild prey with adult females of the other species as intraguild predators. On coffee, a similar set‐up was used, with larvae and adult females of Amblyseius herbicolus Chant and Iphiseiodes zuluagai Denmark & Muma as intraguild prey and intraguild predators, respectively. 4. Domatia on detached, isolated sweet pepper and coffee leaves were either closed with glue or left open, after which larvae and adult predators were released. As a control, larvae were released on leaves with open or closed domatia without an adult predator. 5. Survival of larvae was high in the absence of the adult (intraguild) predator. In the presence of the intraguild predator, survival was significantly higher on leaves with open domatia than on leaves with closed domatia. 6. This shows that even such tiny structures as plant domatia may significantly affect the interaction strength of intraguild predation.     

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.     

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.     

Do domatia mediate mutualistic interactions between coffee plants and predatory mites?

Many plant species possess structures on their leaves that often harbour predatory or fungivorous mites. These so‐called domatia are thought to mediate a mutualistic interaction; the plant gains protection because mites decimate plant pathogenic fungi or herbivores, whereas the mites find shelter in the domatia. We tested this hypothesis using two species of coffee (Coffea spp.) plants that posses domatia consisting of small cavities at the underside of the leaves, and which often harbour mites. We assessed densities of domatia, of the predatory mite Iphiseiodes zuluagai Denmark and Muma (Acari: Phytoseiidae) and of herbivorous mites Oligonychus ilicis (McGregor) (Acari: Tetranychidae) and Brevipalpus phoenicis (Geijskes) (Acari: Tenuipalpidae) on Coffea arabica L. (Rubiaceae) and Coffea canephora Pierre in the field. Over a period of 50 days, C. arabica harboured on average 7.5 times more predatory mites and 0.4–0.66 fewer prey mites than C. canephora. Hence, the higher density of predatory mites on C. arabica could not be explained by higher densities of prey. However, the density of domatia on C. arabica was on average 1.65 times higher than on C. canephora, and within each species, leaves with higher densities of domatia also harboured more predators. This suggests a positive effect of domatia on predatory mites. In the laboratory, survival of adult female predatory mites on leaves of C. arabica with open domatia was indeed significantly higher than on leaves with closed domatia. Hence, predatory mites benefited from the domatia. However, plants with higher densities of domatia did not harbour fewer herbivores. Taken together, our study only provides partial evidence for a mutualistic interaction between coffee plants and predatory mites, mediated by domatia.     

Phytoseiid predator of whitefly feeds on plant tissue

Predatory mites of the family Phytoseiidae feed on herbivorous mites and insects but they also use a variety of non-prey food items, such as pollen and nectar. Plant tissue is another potential food source. We investigated whether plant feeding occurs in the two phytoseiids Euseius scutalis (Athias-Henriot) and Typhlodromips swirskii (Athias-Henriot), which are natural enemies of whiteflies. These predatory mites can suppress populations of Bemisia tabaci (Gennadius) on isolated plants and are candidates for biological control of this pest. Both species can be reared on a diet of pollen, but E. scutalis requires a leaf tissue substrate, suggesting that this species might feed on plant tissue. To test this hypothesis, we applied a systemic insecticide (aldicarb) to cucumber plants and assessed the survival of predatory mites on leaves from insecticide-treated plants and untreated plants, both in presence and absence of pollen. The survival of T. swirskii was not affected by the presence of systemic insecticide in the plant. However, the survival of E. scutalis on leaves from insecticide-treated plants was 10 times lower than on leaves from untreated plants. Since the two species showed similar sensitivity to the insecticide when applied in a slide-dip test, this suggests that E. scutalis ingested insecticide through feeding on the leaf tissue. Mortality on treated leaves was observed both in absence and presence of pollen, suggesting that plant feeding is indispensable for E. scutalis. The extent to which plants are used as food by E. scutalis requires further analysis.     

Attraction of Phytoseiulus persimilis (Acari: Phytoseiidae) towards volatiles from various Tetranychus urticae-infested plant species

Plants infested with the spider mite Tetranychus urticae Koch, may indirectly defend themselves by releasing volatiles that attract the predatory mite Phytoseiulus persimilis Athias-Henriot. Several plants from different plant families that varied in the level of spider mite acceptance were tested in an olfactometer. The predatory mites were significantly attracted to the spider mite-infested leaves of all test plant species. No differences in attractiveness of the infested plant leaves were found for predatory mites reared on spider mites on the different test plants or on lima bean. Thus, experience with the spider mite-induced plant volatiles did not affect the predatory mites. Jasmonic acid was applied to ginkgo leaves to induce a mimic of a spider mite-induced volatile blend, because the spider mites did not survive when incubated on ginkgo. The volatile blend induced in ginkgo by jasmonic acid was slightly attractive to predatory mites. Plants with a high degree of direct defence were thought to invest less in indirect defence than plants with a low degree of direct defence. However, plants that had a strong direct defence such as ginkgo and sweet pepper, did emit induced volatiles that attracted the predatory mite. This indicates that a combination of direct and indirect defence is to some extent compatible in plant species.     

Biological Control of the Bulb Mite, Rhizoglyphus robini , by the Predatory Mite, Hypoaspis aculeifer , on Lilies: Predator-Prey Dynamics in the Soil, under Greenhouse and Field Conditions

We tested the capacity of the soil-dwelling predatory mite, Hypoapsis aculeifer , to control mites attacking lily bulbs. Experiments in the greenhouse and in the field showed that in the absence of predatory mites populations of the bulb mite, Rhizoglyphus robini , on lily bulbs increased, whereas the release of predatory mites either slowed down the increase - as observed in the field - or caused the bulb mites populations to decrease - as observed in the greenhouse. In all cases the population of predatory mites increased as long as bulb mite densities were not too low. However, within the first week after predator release there was usually a sharp decline to 10-40% of the original number released. Greenhouse experiments on intact lily bulbs in pots, boxes and 1 m 2 plots with peat soil showed that when released in a ratio of 1 predator to 2 or 5 prey, the predatory mite, Hypoaspis aculeifer , suppressed populations of bulb mites to less than 10 individual per bulb within 6 weeks. Elimination of bulb mites was observed only when the predator-to-prey ratio at release was equal to 3:1. Field experiments in 2 m 2 plots with intact bulbs in rather compact sandy soil showed that when released in ratio of 1 predator to 1 or 2 prey, the predatory mite, H. aculeifer , did not cause the population of bulb mites to decrease, but it did reduce their population growth. The initial predator-to-prey ratios required to achieve suppression (ca 1:2) or elimination (3:1) in the soil environment are much higher than those required for bulb mite elimination when lily bulb scales were embedded in a medium of vermiculite (ca 1:20). Among the possible causes are: (1) the initial losses of predators in the greenhouse and even more so in the field due to mortality and/or emigration from the experimental plots; (2) the lower temperatures in the greenhouse and especially in the field, which slow down the growth and predation processes and thereby delay prey extinction; and (3) the spatial complexity of the soil environment which creates refuges for the bulb mites.     

Dynamics of refuge use: diurnal, vertical migration by predatory and herbivorous mites within cassava plants

Plants may protect themselves against herbivorous arthropods by providing refuges to predatory arthropods, but they cannot prevent herbivores from taking countermeasures or even from reaping the benefits. To understand whether plants benefit from providing self‐made refuges (so‐called domatia), it is not only necessary to determine the fitness consequences for the plant, but also to assess (1) against which factors the refuge provides protection, (2) why predatory arthropods are more likely to monopolise the refuge, and (3) how herbivorous and predatory arthropods respond to and affect each other in and outside the refuge. In this article, we focus on the last aspect by studying the dynamics of refuge use of a predatory mite (Typhlodromalus aripo) and its consequences for a herbivorous mite (Mononychellus tanajoa) on cassava plants in Benin, West Africa. The refuge, located in‐between the leaf primordia of the cassava apex, is thought to provide protection against abiotic factors and/or intraguild predators. To test whether the predator waits for prey in the apex or comes out, we sampled predator‐prey distributions on leaves and in the apex at 4 hour‐intervals over a period of 24 hours. The predatory mites showed pronounced diurnal changes in within‐plant distribution. They were in the apices during the day, moved to the young leaves during night and returned to the apices the next morning. Nocturnal foraging bouts were more frequent when there were more herbivorous mites on the leaves near the apex. However, the foraging predators elicited an avoidance response by mobile stages of their prey, since these were more abundant on the first 20 leaves below the apex during late afternoon, than on the same leaves during night. These field observations on cassava plants show that (1) during daytime predatory mites monopolise the apical domatia, (2) they forage on young leaves during night and (3) elicit avoidance by within‐plant, vertical migration of mobile stages of the herbivorous mites. We hypothesize that cassava plants benefit from apical domatia by acquiring protection for their photosynthetically most active, young parts, because predatory mites (1) protect primordial leaves in the apex, (2) reduce the densities of herbivorous mites on young leaves, and (3) cause herbivorous mites to move down to less profitable older leaves.     

Phytoseiid predators suppress populations of Bemisia tabaci on cucumber plants with alternative food

Phytoseiids are known to attack whiteflies, but it is an open question whether they can be used for biological control of these pest insects. Preselection experiments in the laboratory showed that two out of five phytoseiid species tested, Euseius scutalis and Typhlodromips swirskii, stood out in terms of their ability to develop and reproduce on a diet of Bemisia tabaci immatures. In this paper, we show that both predators are able to suppress whitefly populations on isolated cucumber plants in a greenhouse. Predatory mites were released 2 weeks in advance of the release of B. tabaci. To enable their survival and promote their population growth, they were provided weekly with alternative food, that is, Typha sp. pollen. A few weeks after whitefly introduction, the numbers of adult whiteflies on plants with predators were consistently lower than on plants without predators, where B. tabaci populations grew exponentially. After 9 weeks, this amounted to a 16- to 21-fold difference in adult whitefly population size. This shows that the two phytoseiid species are promising biocontrol agents of B. tabaci on greenhouse cucumber. This revised version was published online in July 2006 with corrections to the Cover Date.