Host plant manipulation of natural enemies: leaf domatia protect beneficial mites from insect predators

Acarodomatia are small tufts of hair or invaginations in the leaf surface and are frequently inhabited by several taxa of non-plant-feeding mites. For many years, ecologists have hypothesized that these structures represent a mutualistic association between mites and plants where the mites benefit the plant by reducing densities of phytophagous arthropods and epiphytic microorganisms, and domatia benefit the mite by providing protection from stressful environmental conditions, other predaceous arthropods, or both. We tested these hypothesized benefits of domatia to domatia-inhabiting mites in laboratory and growth chamber experiments. In separate experiments we examined whether domatia on the wild grape, Vitis riparia, provided protection against drying humidity conditions or predaceous arthropods to two species of beneficial mite: the mycophagous species Orthotydeus lambi, and the predaceous species Amblyseius andersoni. For both taxa of beneficial mite, domatia significantly increased mite survivorship in the presence of the predatory bug, Orius insidiosus and the coccinellids Coccinella septempunctata and Harmonia varigata. There was no evidence for a protective effect of domatia with a third species of predatory arthropod, lacewing larvae Chrysoperla rufilabris. In contrast, there was no evidence for either species of beneficial mite that domatia provided any protection against low humidity. Thus in this system the primary mechanism by which domatia benefit beneficial mites is by protecting these organisms from other predatory arthropods on the leaf surface.     

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.     

Do leaf domatia mediate a plant–mite mutualism? An experimental test of the effects on predators and herbivores

1. Leaf domatia are tiny structures in leaf vein axils that are widespread among plant taxa and have been described to be typically inhabited by predatory and fungivorous mites. The mutualism hypothesis for the function of leaf domatia predicts that predatory and/or fungivorous mites benefit from having a favourable place to take refuge and reproduce and that plants benefit indirectly from reduced herbivory and/or pathogen attack.
2. The effect of leaf domatia on populations of predatory and herbivorous mites was examined for avocado, Persea americana . In separate experiments, domatia were added to leaves of a variety of avocado plants lacking domatia (Hass) and domatia were blocked on a domatia-bearing variety (Toro Canyon).
3. In two out of the five experiments conducted, domatia-bearing plants had significantly higher numbers of predatory mites compared with controls. Although herbivore numbers were consistently lower on plants with domatia than on plants without domatia, in no case did the presence of leaf domatia result in a statistically significant decrease in herbivorous mite populations.
4. These results suggest that domatia may frequently benefit predatory mites, however, indirect effects on herbivorous mites may not commonly exist or may be too difficult to detect.     

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.     

Domatia reduce larval cannibalism in predatory mites

Abstract.  1. Acarodomatia are small structures on the underside of leaves of many plant species, which are mainly inhabited by carnivorous and fungivorous mites.
2. Domatia are thought to protect these mites against adverse environmental conditions and against predation. They are considered as an indirect plant defence; they provide shelter to predators and fungivores and these in turn protect the plants against herbivores and fungi.
3. We studied the possible role of domatia of coffee ( Coffea arabica L.) (Rubiaceae) and sweet pepper ( Capsicum annum L.) (Solanaceae) in reducing cannibalism in the mites inhabiting the domatia. We measured cannibalism of larvae by adults of the predatory mites Iphiseiodes zuluagai Denmark & Muma and Amblyseius herbicolus Chant on coffee leaf discs and of the predatory mite Iphiseius degenerans (Berl.) on sweet pepper leaf. Domatia were closed with glue or left open.
4. Cannibalism in all three species increased when domatia were closed. With I. degenerans , moreover, we found that the previous diet of the cannibal attenuated the effect of domatia on cannibalism.
5. We conclude that domatia can protect young predatory mites against cannibalism by adults and that the diet of cannibals affects the rate of cannibalism.     

Domatia morphology and mite occupancy of Psychotria horizontalis (Rubiaceae) across the Isthmus of Panama

Leaf domatia are small plant structures in vein axials on the undersides of leaves that are often inhabited by mites of several species. The mites are presumed to benefit the plant because they are predatory or fungivorous. The domatia are thought to provide the mites shelter from predators and changes in relative humidity, and in exchange, the mites protect the plant from small herbivores and fungal spores. Differences in relative humidity can affect food availability, changing the interaction between plants and mites. We examined domatium morphology of the shrub Psychotria horizontalis (Rubiaceae) and its associated mite diversity at three sites along the rainfall gradient of the Isthmus of Panama, during the dry and wet seasons. The dry forest had a domatium morphology consistent with providing greater desiccation protection, with trichomes and a smaller domatium opening relative to domatium size (size/opening ratio). Additionally, this size/opening ratio was significantly higher in the dry season than in the wet season at all three sites. Mite diversity was highest at the intermediate rainfall site with a large degree of overlap with the other sites, whereas the dry site and wet site shared few mite species. More fungivorous mites were present in the moist forests and more facultative feeders on fungal spores and small mites in the dry forest. The average mite size at each site matched the average domatium size at each site. The dry forest had small mites in small domatia, whereas the moist forests had larger mites in larger domatia. While these data are primarily observational, the site and seasonal differences in domatium morphology and mite diversity are consistent with two main hypotheses: (1) that protection from changes in humidity would be particularly important when humidity was low, such as in the dry forest and during the dry season (2) more fungivorous mites would be found in domatia of the moist forests. The data presented here further highlight the close adaptive relationship between leaf domatia on plants and the mites that inhabit them.     

Plants,mites and mutualism: leaf domatia and the abundance and reproduction of mites on Viburnum tinus (Caprifoliaceae)

Associations between mites and leaf domatia have been widely reported, but little is known about their consequences for either plants or mites. By excising domatia from leaves of the laureltinus, Viburnum tinus L. (Caprifoliaceae), in the garden and laboratory, we showed that domatia alter the abundance, distribution, and reproduction of potential plant mutualists. Over 4 months, leaves with domatia on six garden shrubs had 2–36 times more predatory and microbivorous mites, and more mite eggs than leaves without domatia. However, this effect varied among plants and was weaker on one shrub with few mites on its leaves. Domatia also influenced the distribution of mites on leaves. A significantly higher fraction of mites, representing all life stages, was found in vein axils of leaves with domatia than in vein axils on leaves without domatia. Single-leaf experiments in the laboratory showed that domatia enhanced reproduction by the predatory mite, Metaseiulus occidentalis, especially at low relative humidity (30–38%). When domatia were removed, oviposition was reduced significantly only at low relative humidity, suggesting that domatia provide mites with refuge from environmental extremes on the leaf surface. Moreover, the use of domatia by predatory mites may reduce the impact of some plant enemies. In two experiments where prey consumption was measured, M. occidentalis ate significantly higher percentages of the eggs of the two-spotted spider mite (Tetranychus urticae). Our results are consistent with the viewpoint that mite-domatia associations are mutualistic. By directly aiding and abetting the third trophic level, plants with leaf domatia may increase the efficiency of some predaceous and microbivorous mites in consuming plant enemies.     

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.     

Benefits associated with the domatia mediated tritrophic mutualism in the shrub Coprosma lucida

The fitness benefits of plant structural adaptations that increase the effectiveness of fungivores against leaf pathogenic fungi are poorly understood. In a 12‐month field experiment, we investigated the effect of domatia on mite density, the role of these mites in limiting leaf fungi, and the associated effects on plant fitness in the endemic New Zealand shrub, Coprosma lucida. The presence of domatia on mite density was controlled using combinations of domatia blocking, sham blocking, mite addition and mite control using miticide. Limiting access to domatia reduced mite density and increased the proportion of leaves without mites. Mite families represented were predominantly fungivorous/detritivorous (97.2%), and predaceous (2.6%); herbivorous mites were absent. Mites significantly reduced fungal hyphae, fungal spores and pollen, but the effect was surface‐(upper/lower) and density‐dependent with the greatest reduction in fungi occurring over low mite densities. Fungal hyphae reduced leaf longevity, but were associated with increased production of new leaves. Hyphae density on old leaves was negatively correlated with the number of domatia produced on new leaves. New leaves in the mite reduction treatment had slightly reduced levels of carbon but not nitrogen. High levels of fungal infection on the lower surface increased the number of fruit fascicles per shoot, however on the upper surface where fungi were reduced by mites, hyphae density was negatively related to reproduction. The data support a limited interpretation of a fitness benefit for plants with domatia. While domatia increased mite density, control of fungi by mites occurred at lower average densities than supported by plants without functioning domatia. We suggest the primary function of leaf domatia in this mutualism is to increase the probability of a leaf‐level beneficial mite presence rather than to maximise mite density. Many mites are not necessarily better than few mites, but some mites are better than none.     

In search of artificial domatia for predatory mites

Banker plants can enhance biological pest control by providing both floral resources and appropriate oviposition sites, e.g. through acarodomatia, to predator species. The use of materials mimicking domatia i.e. artificial domatia may be an economically favourable alternative to the use of banker plants bearing domatia. The aim of the present study was to identify materials that are able to host eggs of the Neoseiulus californicus predatory mite but not those of the Tetranychus urticae pest mite. In a laboratory experiment, the oviposition of predatory and phytophagous mites were compared in Petri dishes containing leaves. The different modalities compared were (i) natural domatia of Viburnum tinus or (ii) one of twelve potential artificial domatia materials. The overall oviposition response of predatory mites to all artificial domatia was similar to that of the natural domatia. The oviposition of the Tetranychus urticae pest mite did not increase in response to the artificial domatia. Five artificial domatia hosted as many eggs of the predatory mite as observed in the natural domatia. The effect of the physical properties of artificial domatia was also tested and N. californicus was found to favour the artificial domatia that had high heat retention capacities for oviposition. Three of these artificial domatia were tested on rose plants in a greenhouse experiment; none of which enhanced the biological control on the plants under these conditions. The present study highlights the difficulty in identifying and using suitable artificial domatia as substitutes to banker plants in biological pest control efforts.     

Behavioral and population consequences of acarodomatia in grapes on phytoseiid mites (Mesostigmata) and implications for plant breeding

We examined the influence of acarodomatia in the riverbank grape Vitis riparia Michaux (Vitaceae) on the distribution and abundance of predatory mites (Phytoseiidae) and their interactions with herbivorous mites. Acarodomatia are tufts of nonglandular trichomes or pits located in major leaf vein axes of many species of woody perennial plants and are often occupied by predatory and mycophagous mites. In common garden plantings of different accessions of V. riparia we found a significant positive relationship between size of domatia and the abundance of naturally occurring predatory mites. Behavior of adult predatory mites may explain this positive association, in part. In separate laboratory experiments, gravid females of Typhlodromus pyri Scheuten and Amblyseius andersoni Chant spent more time and deposited more eggs on half of a V. riparia leaf with accessible domatia versus the other half in which access to domatia was blocked with pruning tar. Domatia also had population consequences. In an outdoor experiment using potted grapevines, population size of T. pyri and A. andersoni mites was greater on V. riparia with open domatia compared to V. riparia in which domatia were blocked with pruning tar. Population size of predatory mites was also greater on V. riparia with domatia than on Vitis vinifera L., whether their axils were blocked or not. Since V. vinifera have very small domatia, these results indicate that the presence of domatia is important, not just access to vein axils. Elevated predatory mite populations in response to domatia, however, did not translate into differences in the abundance of European red mite Panonychus ulmi (Koch), an important pest of grapes.Overall, these results indicate that domatia in uncultivated V. riparia promote higher densities of some species of generalist phytoseiid mites. However, domatia are small in most cultivated grapes. We crossed females and males of V. riparia that varied in domatia size and reared their offspring and found that average domatia size in the parents was highly correlated (r² = 0.77, slope = 0.55) with average domatia size in offspring (high narrow-sense heritability). Given that V. riparia possesses many other desirable agronomic traits, this result suggests it should be practical to breed for well-developed domatia in cultivated accessions.     

How leaf domatia and induced plant resistance affect herbivores, natural enemies and plant performance

Predators and plant resistance may act together to control herbivorous arthropod populations or antagonistically, which would reduce the control of pest populations. In a field experiment we enhanced predation by adding simulated leaf domatia to plants. Leaf domatia are small structures that often harbor predaceous arthropods that are potentially beneficial to the plant. We also manipulated host plant quality by inducing resistance with controlled, early season exposure of seedlings to spider mite herbivory.
Our manipulations had profound consequences for the natural community of arthropods that inhabited the plants. Leaf domatia had a direct positive effect on abundances of two species of bugs and one species of thrips, all of which are largely predators of herbivores. On leaves with domatia, each of the predators was found inside the domatia two to three times more often than outside the domatia. Eggs of predaceous bugs inside leaf domatia were protected from parasitism compared to eggs outside the domatia. The positive effects of leaf domatia on predator abundances were associated with reduced populations of herbivorous spider mites, aphids, and whiteflies. Plants with experimental leaf domatia showed significantly enhanced reproductive performance.
Induced resistance also affected the community of arthropods. Of the abundant predators, all of which also fed on the plant, only minute pirate bugs were negatively affected by induced resistance. Populations of herbivorous spider mites and whiteflies were directly and negatively affected by induction. In contrast, aphid populations were higher on plants with induced resistance compared to uninduced plants. Effects of induced resistance and domatia were additive for each of the predators and for aphids. However, spider mite and whitefly populations were not suppressed further by employing both induced resistance and domatia compared to each strategy alone. Our manipulations suggest that plant defense strategies can have positive effects on some species and negative effects on others. Negative effects of “resistance traits” on predators and positive effects on some herbivores may reduce the benefits of constitutive expression of resistance traits and may favor inducible defense strategies. Multiple plant strategies such as inducible resistance and morphological traits that aid in the recruitment of predators of herbivores may act together to maximize plant defenses, although they may also be redundant and not act additively.     

Biotic interactions of mites, plants and leaf domatia

Leaf domatia, minute structures that typically house mites and other small arthropods, are produced by an impressive number of plants; however, their role in mediating plant-mite mutualism has only recently been elucidated. New evidence indicates that domatia function primarily as refuges for beneficial mites against predators. The presence of domatia therefore results in more beneficial mites on leaves, fewer pathogen attacks and reduced leaf herbivory. Unexpectedly, herbivorous mites are specialized domatia inhabitants of some plants. By providing refuges for herbivores, however, domatia may stabilize interactions between predator and their mite prey and thereby reduce the chances of herbivore outbreaks. Understanding the ecological mechanisms that promote beneficial interactions between mites and plants could have important implications for pest management.     

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.     

Multiple resource supplements synergistically enhance predatory mite populations

Many plants offer food rewards such as extrafloral nectar and food bodies, which have been shown to attract and retain entomophagous arthropods. In addition to food rewards, plants may possess structures that serve as shelter and/or oviposition sites for beneficial arthropods, so-called domatia. Acarodomatia are commonly used by beneficial mites for oviposition and protection from intraguild predators and adverse climatic conditions (drought). While in nature these food and shelter traits often occur in combination, they have been largely studied in isolation and we know little about how these traits interact, i.e., whether they act independently, antagonistically or synergistically. In the present study, we used citrus seedlings to test the impact of provisioning fibers (as a proxy for acarodomatia), as well as two different categories of food rewards (pollen and sugars) on oviposition and population development of phytoseiid mites. The highest oviposition and abundance of predatory mites was obtained in the treatment where the three resources were offered in combination. The combined impact of the three resources when provided jointly was up to five times higher than the summed impacts of each resource provided individually, thus providing evidence for a three-way synergy between the fibers, pollen and sugars. From an ecological point of view, our results demonstrate that combining multiple indirect defensive traits can strongly enhance the impact on the mutualistic arthropods. Differences in resource provisioning strategies in plant–phytoseiid and plant–ant mutualisms are being discussed. The presented results are of particular importance for our understanding of the functioning of defensive plant-arthropod mutualisms, as well as for the use of predatory mites in conservation- or inundative biological control.     

Oviposition patterns in a predatory mite reduce the risk of egg predation caused by prey

Abstract 1. Predatory arthropods lay their eggs such that their offspring have sufficient prey at their disposal and run a low risk of being eaten by conspecific and heterospecific predators, but what happens if the prey attacks eggs of the predator? 2. The egg distribution and time allocation of adult female predatory mites Iphiseius degenerans as affected by predation of their eggs by prey, the western flower thrips Frankliniella occidentalis, were studied on sweet pepper plants. The predatory mites attack the first instar of thrips but all active stages of thrips are capable of killing the eggs of the predator; however the predatory mite is used for biological control of thrips. 3. The majority of predatory mite eggs was laid on the underside of leaves in hair tufts (domatia). During the experiment, females spent increasing amounts of time in flowers where they fed on pollen and thrips larvae. The risk of predation on predator eggs by thrips was lower on leaves than in flowers where the majority of thrips resides. Moreover, predation risk was higher outside leaf domatia than inside. 4. This suggests that predators avoid ovipositing in places with abundant prey to prevent their eggs from being eaten by thrips.     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

Limited Predator-Induced Dispersal in Whiteflies

Whereas prey are known to avoid habitats with their predators, it is less well established whether they are triggered to emigrate to new habitats when exposed to predators in their current habitat. We studied plant-to-plant dispersal of adult whiteflies in response to the presence of predatory mites on the plant on which the whiteflies were released. These predators attack whitefly eggs and crawlers, but not the adults, which can fly to other plants and can learn to avoid plants with predators. Being tiny and wingless, the predatory mites are slow dispersers compared to adult whiteflies. This offers the whiteflies the opportunity to escape from plants with predatory mites to plants without predators, thus avoiding predation of their offspring. To test for this escape response, a greenhouse experiment was carried out, where whiteflies were released on the first of a row of 5 cucumber plants, 0.6 m or 2 m apart, and predators either on the same plant, on the next plant, or nowhere (control). Adult whiteflies dispersed significantly faster from plants with predatory mites onto neighbouring plants when the plants were 0.6 m apart, but not when plants were 2 m apart. However, the final numbers of whiteflies that had successfully dispersed at the end of the experiments did not differ significantly for either of the two interplant distances. Overall, the proportion of whiteflies that did disperse was low, suggesting that adult whiteflies were apparently reluctant to disperse, even from plants with predators. Our results suggest that this reluctance increases with the distance between the plants, so most likely depends on the uncertainty to find a new plant. Thus, whiteflies do not always venture to fly even when they can easily bridge the distance to another plant.     

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.     

Diapause incidence in the two-spotted spider mite increases due to predator presence, not due to selective predation

We recently reported evidence for increased diapause incidence in the spider mite Tetranychus urticae in presence of the predatory mite Typhlodromus pyri. This effect may arise from (1) selective predation on non-diapause spider mites, (2) predator-induced diapause in spider mites, or (3) both. Using a different strain of T. urticae, we first recovered increased diapause incidence in association with predators. Then, we tested for selective feeding in two-choice experiments with equal numbers of non-diapause and diapause spider mites. We found that the predatory mite had a significant preference for the latter. This indicates that increased diapause incidence in association with predatory mites is not due to selective predation. Therefore, predator-mediated physiological induction of diapause seems a more likely explanation. The cues leading to induction appear to relate to the predators, not their effects, since predation simulated by spider-mite removal or puncturing did not significantly affect diapause incidence. Why spider mites benefit from this response, remains an open question.This revised version was published online in May 2005 with a corrected cover date.