Induced resistance inSolanum dulcamara triggered by the gall miteAceria cladophthirus (Acari: Eriophyoidea)

After infestation ofSolanum dulcamara leaves by the eriophyoid gall miteAceria cladophthirus (Nalepa), induced resistance became operative against subsequent eriophyoid attacks. The protective effect, manifested by reduction in plant damage and/or mite proliferation, lasted up to 40 days. When the challenger wasA. cladophthirus, the number and size of lesions decreased significantly but mite mortality was not enhanced. When the rust miteThamnacus solani was used as challenger, both russeting symptoms and mite survival rate were strongly reduced. AsA. cladophthirus-induced resistance is relatively permanent, it opens new prospects for eriophyoid control in the field.     

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

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

The gall miteAceria cladophthirus. I. Life-cycle,survival outside the gall and symptoms' expression on susceptible or resistantSolanum dulcamara plants

The gall miteAceria cladophthirus (Nalepa) is able to survice outside its gall on detached leaves ofSolanum dulcamara L. kept under non-aseptic in-vitro conditions. The survival rate of the females on susceptible leaves is about 90% after 1 day and 85% for the following days. In contrast, on resistant leaves, less than 40$ survive after 1 day while necrotic local lesions develop and later the mortality increases severely. However, the mite only completes its life-cycle on susceptible leaves. The life-cycle forA. cladophthirus takes about 12 days: six days for egg incubation and six days for two instars growth. Its life-history is simple, without alternating females specialised for hibernation; arrhenotokous parthenogenesis occurs in experimental conditions. Eggs do not play any role in gall formation. Immature stages induce gall symptoms but are less efficient than females. Mite feeding only induces complete gall symptoms on the less differentiated leaves of susceptible shoots. On susceptible detached leaves, gall symptoms are similar but weaker and their intensity decreases with increasing leaf age; fully expanded leaves remain free of symptoms.On resistant plants, mite feeding induces a hypersensitive response: necrotic local lesions, about 350 m in diameter, appear both on shoots and on detached leaves. However, young leaves develop smaller lesions than old ones. Females induce larger lesions than first larvae. Once necrosis formation is initiated, it proceeds to completion regardless of the duration of mite feeding.     

Factors influencing the acaricidal activity of flucycloxuron

Contact activities of flucycloxuron on immature stages of the two-spotted spider mite (Tetranychus urticae (Koch)) and the European red mite (Panonychus ulmi (Koch)) gradually decrease in the successive developmental stages. The levels of contact activity of flucycloxuron on larvae and protonymphs ofT. urticae andP. ulmi are of the same order. Deutonymphs ofT. urticae are less susceptible to contact activity than the similar stage ofP. ulmi. In adultT. urticae, the transovarial ovicidal activity was used as an indicator for cuticular penetration. More than 90% of the maximal penetration into adult mites occurs within 8 h. Reversibility of the transovarial activity was not observed after 24 h, but did occur after a subsequent 48 h stay on untreated leaves. The ovo-larvicidal activity of flucycloxuron onP. ulmi after treatment of apple leaves is strongly negatively influenced by leaf age, partly by lower retention of the spray liquid on the leaves. Leaf penetration was measured by application of flucycloxuron on leaf uppersides and assessment of the transovarial activity in mites (P. ulmi orT. urticae) infested on the undersides, one day after treatment. In this test system, leaf penetration was found to be strongly species dependent. Penetration was high in cucumber, moderate in French beans, cotton, roses and strawberry, but low in apple and pepper. Leaf penetration in French bean plants is drastically reduced at increasing leaf age. The overall positive effect of increase in relative air humidity on leaf penetration, is statistically highly significant (P=0.001) for French beans and almost significant (P=0.08) for cucumbers. WithT. urticae on French bean it was found that in this test flucycloxuron needs more than one day for maximal leaf penetration. Although in apple leaves penetration from uppersides was low, penetration from undersides was much higher. The surfactants Arkopal N 130, Silwet L-77 and X2-5309 enhance penetration from leaf under-sides.     

Prey preference of the phytoseiid miteTyphlodromus pyri 1. Response to volatile kairomones

Using a Y-tube olfactometer, a study has been made of the response of females of the predatory miteTyphlodromus pyri Scheuten (Acarina: Phytoseiidae) to volatile kairomones of three prey species: the European red spider mite (Panonychus ulmi (Koch)), the two-spotted spider mite (Tetranychus urticae Koch) and the apple rust mite (Aculus schlechtendali (Nalepa)).Predators that had been reared onT. urticae responded only to the volatile kairomone ofP. ulmi. In contrast, when reared onVicia faba L. pollen, they responded to the kairomones of all three prey species. Pollen-reared predators, offered a choice between kairomones of two different prey species, prefer theP. ulmi kairomone to those ofA. schlechtendali orT. urticae.The difference in response between predators reared onV. faba pollen andT. urticae seems to be caused by the low carotenoid content ofV. faba pollen. Predators that had been reared onV. faba pollen mixed with crystalline -carotene behaved in a way similar to conspecific that had been reared on the carotenoid-rich prey miteT. urticae. Obviously, pollen-rearedT. pyri females are in need of carotenoids, which can be obtained from, e.g.,P. ulmi, T. urticae orA. schlechtendali. This may explain why pollen-reared predators respond to more prey species thanT. urticae-reared predators.WhyT. pyri females need carotenoids has not been established. The only known function of carotenoids in mites is involvement in diapause induction. However, as pollen-rearedT. pyri enter reproductive diapause under short-day conditions, they either extract sufficient amounts of carotenoids fromV. faba pollen, or do not need carotenoids for diapause induction.Apart from the effect of dietary requirements on prey selection, food deprivation also affects the predator's response to kairomones. All the data mentioned above have been obtained for predators that had been starved for 20 h. Predators that had been reared onT. urticae and starved for 48 h before the experiment did respond to the volatile kairomone ofT. urticae in contrast to predators from the same culture that had been starved for 20 h. Thus foraging decisions byT. pyri are affected by both starvation time and specific hunger for carotenoids.     

Diapause induction and duration in the phytoseiid miteTyphlodromus pyri

A photoperiod of 8L/16D for two weeks was used to distinguish between diapausing and non-diapausingTyphlodromus pyri Scheuten. A diet ofPanonychus ulmi orTetranychus urticae, or pollen ofVicia faba did not influence preovipositional periods of diapausing mites. In mid-September, 88% ofT. pyri collected from an insectary were in diapause. The critical day-length appeared to be between 12.5 and 13.5 h. Diapause duration was greatest in mites collected in September–November, becoming progressively less in mites collected later in the winter. By mid-April, ca. 50% of mites collected from the orchard and insectary oviposited promptly when cultured in the laboratory.Typhlodromus pyri eggs and larvae were present on leaves in early May. At this time, only 4% ofP. ulmi winter eggs had hatched. Diapause terminated most quickly inT. pyri kept in an 18L/6D photoperiod, followed by 24L/0D and 0L/24D. Longest preoviposition periods were recorded for mites kept in 8L/16D photoperiods.     

The presence of webbing affects the oviposition rate of two-spotted spider mites, <Emphasis Type="Italic">Tetranychus urticae</Emphasis> (Acari: Tetranychidae)

Several species of tetranychid mites including Tetranychus urticae Koch (Acari: Tetranychidae) construct complicated three-dimensional webs on plant leaves. These webs provide protection against biotic and abiotic stress. As producing web is likely to entail a cost, mites that arrive on a leaf with web are expected to refrain from producing it, because they will gain the benefit of protection from the existing web. Mites that produce less web may then allocate resources that are not spent on web construction to other fitness-enhancing activities, such as laying eggs. To test this, the oviposition rate of T. urticae adult females was examined on leaves with web. As a control, we used leaves where the web had been removed, hence both types of leaves had been exposed to conspecifics previously and were thus damaged. On leaves with web, the oviposition rate of T. urticae females was higher than on leaves where the web had been removed. Therefore, the presence of web constructed by conspecifics enhanced the oviposition rate of T. urticae females. This provides indirect evidence that mites use the web constructed by conspecifics and thereby save resources that can be allocated to other traits that enhance reproductive success.     

Herbivore performance and plant defense after sequential attacks by inducing and suppressing herbivores

It is well known that herbivore-induced plant defenses alter host plant quality and can affect the behavior and performance of later arriving herbivores.Effects of sequential attacks by herbivores that either suppress or induce plant defenses are less well studied.We sequentially infested leaves of tomato plants with a strain of the phytophagous spider mite Tetranychus urticae that induces plant defenses and the closely related Tetranychus evansi, which suppresses plant defenses.Plant quality was quantified through oviposifion of both spider mite species and by measuring proteinase inhibitor activity using plant material that had been sequentially attacked by both herbivore species.Spider-mite oviposifion data show that T.evansi could suppress an earlier induction of plant defenses by T.urticae,and T.urticae could induce defenses in plants previously attacked by T.evansi in 1 day.Longer attacks by the second species did not result in further changes in oviposifion.Proteinase inhibitor activity levels showed that T.evansi suppressed the high activity levels induced by T.urticae to constitutive levels in 1 day,and further suppressed activity to levels similar to those in plants attacked by T.evansi alone.Attacks by T.urticae induced proteinase inhibitor activity in plants previously attacked by T.evansi,eventually to similar levels as induced by T.urticae alone.Hence,plant quality and plant defenses were significantly affected by sequential attacks and the order of attack does not affect subsequent performance,but does affect proteinase inhibitor activity levels.Based on our results,we discuss the evolution of suppression of plant defenses.     

Long-term stability in the interaction betweenTetranychus urticae andPhytoseiulus persimilis producing successful integrated control on roses in southeast Queensland

Phytoseiulus persimilis Athias-Henriot successfully controlledTetranychus urticae Koch on rose hedges in an integrated mite control programme in two unheated plastic-covered tunnels at Cleveland in southern Queensland. Mite populations were monitored fortnightly for nearly three years in one tunnel and for two years in a second. Once established,P. persimilis quickly suppressedT. urticae and then persisted. A relatively stable, long-term, low-level interaction developed between the two species under normal commercial conditions. In one tunnel, the production (upper) level of the four rose hedges contained fewer mites of both species per compound leaf (1.71 of all stages ofT. urticae, 0.38 motiles; 0.25 of all stages ofP. persimilis, 0.12 motiles) than the maintenance (lower) layer (2.02 of all stages ofT. urticae, 0.75 motiles; 0.35 of all stages ofP. persimilis, 0.21 motiles). On 68% of sampling occasions, 10% or less of compound leaves in the production layer were infested withT. urticae (all stages including eggs) and control was excellent throughout the 30-month period afterP. persimilis was established, there being no economic losses. In the second tunnel, there was no significant difference between mite numbers in the production and maintenance layers of the two hedges examined. The overall mean number of all stages ofT. urticae per compound leaf was 3.2 (1.45 motiles), and ofP. persimilis 0.46 (0.25 motiles). Control here was slightly less effective than in the first tunnel, but was still satisfactory, with 10% or less of leaves being infested on 45.5% of sampling occasions (20% or less on 66% of sampling occasions) withT. urticae (all stages including eggs).Phytoseiulus persimilis was not reintroduced or redistributed during the course of the observations. An auxiliary miticide (clofentezine) was applied on only one occasion to selected areas in the second tunnel. The influence of high temperatures on control byP. persimilis is 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.     

Evolutionary adaptation to host plants in a laboratory population of the phytophagous mite Tetranychus urticae Koch

Summary For evolutionary expansion of host range to occur in an herbivore population, genetic variation in ability to survive on and/or accept new hosts must be present. To determine whether a population of the phytophagous mite Tetranychus urticae contained such variation, I established lines from the population on two hosts on which mites initially showed both high juvenile mortality and low acceptance, tomato and broccoli. In less than ten generations, mites from the line kept on each host showed both lower mortality and greater acceptance on it than mites from a control line kept on lima bean, a favorable host for T. urticae. Host acceptance was measured by the proportion of mites attempting to disperse from leaves of the host. The line kept on tomato but not the one kept on broccoli also increased in development rate on its host. These results and those of a similar previous experiment on cucumber indicate that T. urticae populations can adapt to a diversity of initially unfavorable hosts. T. urticae populations therefore should be able to respond to temporal and spatial variation in host availability by adapting to the most abundant hosts.     

Environmental actions of agrochemicals 1. Side-effects of the herbicide 3-amino-1,2,4-triazole on a laboratory acarine/host-plant interaction (Tetranychus urticae/Phaseolus vulgaris) as revealed by electron microscopy

Foliar and soil application in concentrations below the recommended rate of the herbicide 3-amino-1,2,4-triazole to the host plantPhaseolus vulgaris L. results in structural alterations of the protein-synthesizing apparatus of midgut and salivary-gland cells of the phytophagous spider miteTetranychus urticae Koch (Acari: Tetranychidae) independent of its mode of application. With prolonged incubation times cytological defects become more intense, and spread to more cells and tissues. Resultant effects on yolk and egg formation were expressed as an inhibition of egg deposition that led to a decrease in the reproduction rate ofT. urticae.Consequences of 3-amino-1,2,4-triazole action onT. urticae are discussed with regard to its value to a host-plant/parasite model, agricultural practices and environmental impacts.     

Induced feeding preferences in dicofol-resistant two-spotted spider mites (Acari: Tetranychidae)

Preadult rearing conditions affected the behavior of dicofol-resistant two-spotted spider mites (Tetranychus urticae). Resistant spider mites reared on dicofol-treated leaves initiated a significantly greater number of feeding bouts on dicofol-treated leaves than did genetically identical spider mites reared on residue-free leaves. Therefore the prior exposure of resistant spider mites resulted in induced feeding preferences that could exacerbate the potential outcome of the resistance by resulting in greater amounts of feeding by resistant individuals on dicofol-treated areas. Since resistant individuals that had not experienced dicofol in their lifetime did not display this feeding preference, avoidance of this phenomenon of induced feeding preference may be an undescribed value of rotations of pesticides.     

Mitochondrial DNA from a spider mite: isolation,restriction map and partial sequence of the cytochrome oxidase subunit I gene

Mitochondrial (mt) DNA of the phytophagous miteTetranychus urticae was purified and a restriction map was constructed. The 12.5 kb long genome is the shortest animal mtDNA known. A 564 bp clone comprising part of the gene for cytochrome oxidase subunit I was sequenced. As has been found in insects, the mitochondrial sequences of mites are extremely A+T rich (75% on average, 96.5% at the third codon position).     

A key volatile infochemical that elicits a strong olfactory response of the predatory mite <Emphasis Type="Italic">Neoseiulus californicus</Emphasis>, an important natural enemy of the two-spotted spider mite <Emphasis Type="Italic">Tetranychus urticae</Emphasis>

Herbivore-induced plant volatiles (HIPVs) emitted from lima bean leaves infested with the two-spotted spider mites Tetranychus urticae strongly attract the predatory mites Neoseiulus californicus. Among these HIPVs, methyl salicylate and linalool can attract the predators. Three green-leaf volatiles (GLVs) of (Z)-3-hexen-1-ol, (Z)-3-hexenyl acetate and (E)-2-hexenal, found in the odor blends from T. urticae-infested leaves and physically damaged leaves, can also attract the predators. To search for a strong predator attractant, the olfactory responses of N. californicus to each synthetic compound or their combinations were investigated in a Y-tube olfactometer. When presented a choice between a mixture of the five compounds (i.e. the two HIPVs and the three GLVs) and T. urticae-infested leaves, N. californicus did not discriminate between these odor sources. The same trend was observed when either a mixture of the two HIPVs or methyl salicylate vs. T. urticae-infested leaves were compared. In contrast, the predators preferred T. urticae-infested leaves to linalool, each of the three GLVs, or a mixture of the three GLVs. These results indicated that methyl salicylate is a strong predator attractant, and its potential attractiveness almost equaled that of the blend of HIPVs from T. urticae-infested leaves.     

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.     

Voluntary Falling in Spider Mites in Response to Different Ecological Conditions at Landing Points

We examined voluntary-falling behaviour by adult females of the two-spotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae) and one of its major predators Neoseiulus californicus McGregor (Acari: Phytoseiidae). Experiments were conducted using a setup in which mites could only move onto one of two landing points by falling. Significantly more T. urticae females fell onto available food leaves compared to non-food or heavily infested leaves, whereas significantly fewer females fell onto leaves with the predatory mite N. californicus compared to leaves without the predator. This suggests that spider mites can actively choose on which patch to land on the basis of food quality and predation risk on the patch. Using the same experimental setup, starved N. californicus females never fell, suggesting that falling T. urticae females gain the potential advantage of predator avoidance.     

Amensalism via webs causes unidirectional shifts of dominance in spider mite communities

Competitive displacement is considered the most severe consequence of interspecific competition; if a superior competitor invades the habitat of an inferior species, the inferior species will be displaced. Most displacements previously reported among arthropods were caused by exotic species. The lack of investigation of displacement among native species may be due to their apparently harmonious coexistence, even if it is equivalent to an outcome of interspecific association. A seasonal change in the species composition of spider mites, from Panonychus ulmi to Tetranychus urticae, is observed in apple trees worldwide. Previous laboratory experiments have revealed amensal effects of T. urticae on P. ulmi via their webs. Using manipulation experiments in an orchard, we tested whether this seasonal change in species composition occurred as the result of interspecific competition between these spider mites. Invasion by T. urticae prevented an increase in P. ulmi densities throughout the experimental periods. Degree of overlap relative to the independent distribution on a leaf-surface basis (ω _S) changed from positive to negative with increasing density of T. urticae. T. urticae invasion drove P. ulmi toward upper leaf surfaces (competitor-free space). The niche adjustment by P. ulmi occurred between leaf surfaces but not among leaves. Our findings show that asymmetrical competition between T. urticae and P. ulmi plays an important role in this unidirectional displacement and that the existence of refuges within a leaf produces the apparently harmonious coexistence of the mites and obscures their negative association.     

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.     

Integrated control of two spotted spider mite,Tetranychus urticae on cucumber grown under plastichouse conditions in egypt

Microclimatic conditions were monitored in cucumber grown under unheated plastichouses in three consecutive years, i.e. from 1990 to 1992 in Egypt. The relative humidity in the plastichouses constructed on either sandy or clay soil did not exceed 30% during mid-day time, while the average temperature did not drop below 30°C. Such conditions favoured population build up of the two spotted spider miteTetranychus urticae Koch. In 1993 and 1994, three plastichouses planted to cucumber, cultivar Rawa or Hana were selected. Relative humidity in two of them was maintained over 50% during the day using spraying nozzles, the third being kept as control. Mildew diseases induced by increased leaf wetness needed to be managed using fungicides.Phyioseiulus persimilis andP. longipes were released in the first two plastichouses, while acaricides were applied in the control. In early MayT. urticae population increase on the growing top of cucumber plants required two acaricidal applications as predatory mites did not migrate to the apical leaves which were too warm and dry. More than 80% of the maximum yield was obtained on average in plastichouses using increased relative humidity, but without any acaricidal applications.