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.     

Efficacy of Metarhizium anisopliae in controlling the two‐spotted spider mite Tetranychus urticae on common bean in screenhouse and field experiments

The efficacy of aqueous and emulsifiable formulations of the fungus Metarhizium anisopliae isolate ICIPE78 was evaluated on the population density of Tetranychus urticae infesting common bean plants under screenhouse and field conditions. Synthetic acaricide abamectin was included as a check. Bean plants were artificially infested with T. urticae and allowed to multiply. Three treatments were applied in the screenhouse and 1 treatment in field trials. Mite density was recorded 2 d before spraying and weekly postspraying. The number of pods per plant, number of seeds per pod, and the dry weight of seeds per plant were recorded only in the screenhouse trials. In both screenhouse and field trials, fungal formulations applied at the concentration of 10⁸ conidia/mL and the acaricide reduced the population density of mites as compared to the controls. There were significant differences in T. urticae population densities between the treatments at the various post‐spraying sampling dates. In the screenhouse, the mite densities were near zero from 3‐week postspraying in the treated leaves. At 4‐week postspraying, there were no more leaves in the untreated control (T1) and in the control water + Silwet‐L77 (T2). Fungal formulations were as effective as abamectin in reducing mite densities in both screenhouse and field experiments. There were significant differences in the production parameters during the 2 screenhouse trials, with fungal and abamectin treatments generally having the highest yield. Results of this study underline the potential of the M. anisopliae isolate ICIPE78 as an alternative to acaricides for T. urticae management.     

Evaluation of a damage threshold for two-spotted spider mites, Tetranychus urticae Koch (Acari: Tetranychidae), in hop culture

Damage caused by two‐spotted spider mites (Tetranychus urticae) at harvest to yield, quality (measured in percentage α‐acids content) and cone infestation was assessed on hop cvs Hallertauer Magnum, Hallertauer Tradition and Perle. Acaricide‐untreated hop plants with known levels of T. urticae infestation were compared with neighbouring acaricide‐treated plants. Although in 24 of the 36 experimental harvests the untreated hop plants had spider mite infestations of > 100 mites leaf^?1, yields and α‐acids content from the untreated plants were significantly lower than the treated plants in only four instances. However, although mite infestation of cones from untreated hops were significantly higher than acaricide‐treated plants in 27 of the 36 cases, in only one instance did that cause economic loss. Spider mite infestation levels of c. 90 mites leaf^?1 are tolerable at harvest time with little or no risk of causing economic loss to hop growers.     

The effect of pyrethroid lambdacyhalothrin applications on the spatial distribution of phytophagous and predatory mites in apple orchards

The spatial distribution of three phytophagous mites,Panonychus ulmi (Koch),Tetranychus urticae Koch andAculus schlechtendali (Nalepa), and two predacious mites,Zetzellia mali (Ewing) andAmblyseius fallacis (Garman), and the effect of pyrethroid lambdacyhalothrin applications on mite spatial dispersion were investigated over a 3-year period in an apple orchard in Ontario. The index of dispersion and the slope of Taylor's power law were used to evaluate dispersion patterns of mites. Panonychus ulmi showed that between-tree spatial variation decreased with an increase of population densities, whereas between-leaf variation increased with population densities. With all other four species it appeared that between-tree variation is much greater than between-leaf variation at all field population density levels. The values ofb by Taylor's power law suggested that all five species of mites are aggregated, but that in generalP. ulmi andT. urticae (b=1.427–1.872) are more aggregated than their predators (b=1.254–1.393). Taylor's regression technique suggests that pyrethroid applications causedP. ulmi, T. urticae, Z. mali andA. fallacis to be less aggregated whileA. schlechtendali was more aggregated. The impact of changes in mite spatial distribution following pyrethroid applications on sampling plans is discussed.     

The effect of groundcover and herbicide treatment on twospotted spider mite density and dispersal in southern Oregon pear orchards

A total of 49 groundcover plant species representing 47 genera in 22 families were identified from a survey of 5 pear orchards. Density of twospotted spider mite (Tetranychus urticae Koch) inhabiting these plants was estimated visually several times during the field season. Plants were ranked in 1 of 3 categories depending on mite densities found on these plants during the summer. T. urticae was found to be highly abundant (category 3) on 26 species, at lower densities on 10 species (category 2), and was rarely or never found on the remaining 12 species (category 1). Dispersal of mites from groundcover plants into trees was found to be highly variable within and between orchards. Within orchard dispersal appeared to be related to the distribution and abundance of category 3 host plants in the orchard. Variability between orchards may also be affected by groundcover management techniques and levels of acaricide resistance in T. urticae. The use of herbicides to control groundcover plants significantly increased the dispersal of T. urticae into the orchard trees.     

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.     

Allocating time between feeding, resting and moving by the two-spotted spider mite, Tetranychus urticae and its predator Phytoseiulus persimilis

This study characterizes the timing of feeding, moving and resting for the two-spotted spider mite, Tetranychus urticae Koch and a phytoseiid predator, Phytoseiulus persimilis Athias-Henriot. Feeding is the interaction between T. urticae and plants, and between P. persimilis and T. urticae. Movement plays a key role in locating new food resources. Both activities are closely related to survival and reproduction. We measured the time allocated to these behaviours at four ages of the spider mite (juveniles, adult females immediately after moult and adult females 1 and 3 days after moult) and two ages of the predatory mite (juveniles and adult females). We also examined the effect of previous spider mite-inflicted leaf damage on the spider mite behaviour. Juveniles of both the spider mite and the predatory mite moved around less than their adult counterparts. Newly emerged adult female spider mites spent most of their time moving, stopping only to feed. This represents the teneral phase, during which adult female spider mites are most likely to disperse. With the exception of this age group, spider mites moved more and fed less on previously damaged than on clean leaves. Because of this, the spider mite behaviour was initially more variable on damaged leaves. Phytoseiulus persimilis rested at all stages for a much larger percentage of the time and spent less time feeding than did T. urticae; the predators invariably rested in close proximity to the prey. Compared to adult predators, juveniles spent approximately four times as long handling a prey egg. The predator-prey interaction is dependent upon the local movement of both the predators and prey. These details of individual behaviours in a multispecies environment can provide an understanding of population dynamics.     

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

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

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

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

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

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

The presence of <Emphasis Type="Italic">Tetranychus urticae</Emphasis> (Acari: Tetranychidae) and its predators on plants in the ground cover in commercially treated vineyards

The presence of Tetranychus urticae and its predators on plants in the ground cover in commercially treated vineyards in the Hex River Valley, Western Cape Province, South Africa was investigated. Six Barlinka and six Dauphine vineyards were surveyed. Leaf samples from the plants on the vineyard floor were taken at monthly intervals and microscopically examined for spider mites and their predators. The coverage of the plants was determined using a co-ordinate sampling system. A wide variety of plant species was found on the vineyard floor. Most of the spider mites found on these plants were T. urticae. The phytoseiid mites on the plants were Euseius rubicolus, Neoseiulus californicus and an undescribed Typhlodromus species. The presence of these plants made it possible for T. urticae and phytoseiid mites to occur throughout the year, with a similar pattern in their seasonal activity on the vineyard floor and the vine leaves.     

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

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

Expression of a barley cystatin gene in maize enhances resistance against phytophagous mites by altering their cysteine-proteases

Phytocystatins are inhibitors of cysteine-proteases from plants putatively involved in plant defence based on their capability of inhibit heterologous enzymes. We have previously characterised the whole cystatin gene family members from barley (HvCPI-1 to HvCPI-13). The aim of this study was to assess the effects of barley cystatins on two phytophagous spider mites, Tetranychus urticae and Brevipalpus chilensis. The determination of proteolytic activity profile in both mite species showed the presence of the cysteine-proteases, putative targets of cystatins, among other enzymatic activities. All barley cystatins, except HvCPI-1 and HvCPI-7, inhibited in vitro mite cathepsin L- and/or cathepsin B-like activities, HvCPI-6 being the strongest inhibitor for both mite species. Transgenic maize plants expressing HvCPI-6 protein were generated and the functional integrity of the cystatin transgene was confirmed by in vitro inhibitory effect observed against T. urticae and B. chilensis protein extracts. Feeding experiments impaired on transgenic lines performed with T. urticae impaired mite development and reproductive performance. Besides, a significant reduction of cathepsin L-like and/or cathepsin B-like activities was observed when the spider mite fed on maize plants expressing HvCPI-6 cystatin. These findings reveal the potential of barley cystatins as acaricide proteins to protect plants against two important mite pests.     

Survey of natural enemies of spider mites (Acari: Tetranychidae) in citrus orchards in eastern Spain

Field surveys were conducted from 2004 to 2007 to determine the species composition and relative abundance of natural enemies associated with colonies of either the citrus red mite, Panonychus citri, or the two spotted spider mite, Tetranychus urticae, in Valencian citrus orchards (eastern Spain). Fourteen species were recorded, six phytoseiid mites and eight insect predators. Two of them are reported for the first time on citrus in Spain and two more are first reports as predators associated with T. urticae. The community of predators associated with T. urticae and P. citri was almost identical, and the Morisita–Horn index of similarity between both natural enemy complexes was close to one, suggesting that predators forage on both pest species. Quantifying the presence of many known spider mites predators in Valencian citrus orchards is an important first step towards spider mite control. A challenge for future studies will be to establish conservation and/or augmentation management strategies for these predators, especially to improve T. urticae biological control.     

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.     

Environmental conditions affecting the strength of induced resistance against mites in cotton

Summary Spider mites (Tetranychus urticae) were raised on cotton plants that had been damaged by a previous bout of mite feeding (T. turkestani) and on control plants that were previously not exposed to mites. The effect of induced plant responses upon mite populations was variable ranging from a 4-fold reduction in population growth to no difference at all. The strength of induced resistance was greatest when the population growth of mites was low for other, unknown, reasons. When mite population growth on control plants was great, the effects of induced resistance were diminished.Mite population growth was inversely related to levels of initial damage caused by previous feeding. There was no evidence of a damage threshold that needed to be exceeded before induced resistance became effective. Increased levels of initial damage were not associated with morphological changes in the plants.     

The distribution of herbivores between leaves matches their performance only in the absence of competitors

Few studies have tested how plant quality and the presence of competitors interact in determining how herbivores choose between different leaves within a plant. We investigated this in two herbivorous spider mites sharing tomato plants: Tetranychus urticae, which generally induces plant defenses, and Tetranychus evansi, which suppresses them, creating asymmetrical effects on coinfesting competitors. On uninfested plants, both herbivore species preferred young leaves, coinciding with increased mite performance. On plants with heterospecifics, the mites did not prefer leaves on which they had a better performance. In particular, T. urticae avoided leaves infested with T. evansi, which is in agreement with T. urticae being outcompeted by T. evansi. In contrast, T. evansi did not avoid leaves with the other species, but distributed itself evenly over plants infested with heterospecifics. We hypothesize that this behavior of T. evansi may prevent further spread of T. urticae over the shared plant. Our results indicate that leaf age determines within‐plant distribution of herbivores only in absence of competitors. Moreover, they show that this distribution depends on the order of arrival of competitors and on their effects on each other, with herbivores showing differences in behavior within the plant as a possible response to the outcome of those interactions.     

Predation by Allothrombium pulvinum on the spider mites Tetranychus urticae and Amphitetranychus viennensis: Predation Rate, Prey Preference and Functional Response

The deutonymphs of Allothrombium pulvinum Ewing (Acari: Trombidiidae) are among the most important natural enemies of spider mites in North, North East and West Iran. In this study, maximum predation rate and preference experiments were conducted with A. pulvinum deutonymphs on apple leaf discs, to determine their preference for either of two spider mite species: Amphitetranychus viennensis (Zacher) and Tetranychus urticae Koch (Acari: Tetranychidae). Maximum predation rate tests showed that the predatory mite consumed more eggs and females of T. urticae than of A. viennensis. Furthermore, the Manly’s preference index for eggs and females of T. urticae confirmed that T. urticae were the preferred prey. The functional response of A. pulvinum deutonymphs on females of T. urticae was examined over a 24-h period. Predation of A. pulvinum deutonymphs presented with females of T. urticae followed a type III functional response. Estimated handling time for the predatory mites was 4.51 h and attack coefficient b, which describes the changes in attack rate with prey densities in a type III functional response, was 0.021.     

The spatial and temporal distribution of predatory and phytophagous mites in field-grown strawberry in the UK

Extensive sampling of strawberry plants in everbearing and June-bearing strawberry plantations and on potted plants showed that different species of mites were spatially separated. Of the two phytophagous species recorded, Tetranychus urticae was most abundant on old leaves and Phytonemus pallidus on folded leaves and flower/fruit clusters. Predatory phytoseiid mites were found on all plant parts but different species were spatially separated; Neoseiulus cucumeris and N. aurescens were found mostly on folded leaves and clusters, and N. californicus and Phytoseiulus persimilis on old and medium aged leaves. No Typhlodromus pyri were found in the field plantations. These patterns of distribution did not change over sampling dates in summer and early autumn. An understanding of this within-plant zonation of mite species is important when studying predator–prey interactions and when designing sampling strategies for strawberry. A programme to sample the entire mite system on strawberry should be stratified to include all the above mentioned parts of the plant. Different sampling protocols, as appropriate, are required for sampling different pest species and their associated predators.