Possible ecological consequences of heterospecific mating behavior in two tetranychid mites

Interspecific mating between the two-spotted spider mite,Tetranychus urticae Koch, and the Banks grass mite,Oligonychus pratensis (Banks), was documented using laboratory populations. The incidence of mating betweenT. urticae males andO. pratensis females was 26.0%, while that for the reciprocal mating was 18.8%. The incidence of mating was affected by both male and female species. Such matings may have several important ecological consequences. Interspecific matings resulted in all-male progenies. Thus, progeny sex ratios may be distorted by misdirected mating behavior. In addition, heterospecific mating resulted in lower fecundity than conspecific matings in the two-spotted spider mite, although not in the Banks grass mite. Aerial dispersal behavior of the two-spotted spider mite was also affected. Under crowded conditions and deteriorating resource quality, female mites exhibit an aerial dispersal posture that helps them to become airborne, and allows them to disperse long distances. Forty-two percent ofT. urticae females that mated with conspecific males exhibited this dispersal behavior, compared to only 3.6% for virgin females. The incidence of aerial dispersal behavior for females that mated with heterospecific males was intermediate (27.3%). The effects of these behavioral alterations on male and female fitness may depend on the population structure and resource distribution.     

Synomone-induced suppression of take-off in the phytoseiid mite Phytoseiulus persimilis Athias-Henriot

Plant-inhabiting predatory mites in the family Phytoseiidae are known to disperse passively on air currents. In this article ww analyse observations on the behaviour that initiates aerial dispersal, the so-called take-off behaviour. When starved for 24 hours at 25°C and 35% RH, about 80% of the females of Phytoseiulus persimilis Athias-Henriot became airborne during 10 minute exposure to wind velocities of 2 ms^-1 or higher. However, take-off was suppressed when females were exposed to volatile chemicals emanating from leaves that had been infested by two-spotted spider mites (Tetranychus urticae Koch) during one day preceding the experiments. This result is the first unambiguous proof that phytoseiid mites exert control over take-off. Interestingly, the females of the predator strain under study did not show the characteristic upright posture that was hypothesized to be important for take-off in two other species of phytoseiid mites (Amblyseius fallacis Garman and Metaseiulus occidentalis (Nesbitt)). These observations shed new light on the behaviour involved in controlling take-off. It is suggested that take-off control is exerted mainly via the grasp of the claws and the adhesive empodia in a way reminiscent of that described for aphids.     

Influence of dispersal from almonds on the population dynamics and acaricide resistance frequencies of spider mites infesting neighboring cotton

Neighboring almond and cotton fields were sampled for spider mites in four locations in the San Joaquin Valley of California. The dominant species in the almonds wasTetranychus pacificus McGregor. In three cotton sites.T. pacificus was present in significantly higher densities near the almonds on at least one sampling date. In contrast.T. urticae Koch andT. turkestani Ugarov & Nikolski were equally abundant across the cotton fields. Almonds appeared to act as a continuous early-season source ofT. pacificus for cotton, with peaks in aerial dispersal from almonds occurring due to overcrowding, plant water stress, and applications of repellent acaricides. Cotton, which experienced little water stress, supported very high densities of spider mites and so acted primarily as a sink for spider-mite dispersal from almonds and other field crops throughout the growth-season. The frequencies of resistance expressed byT. pacificus andT. urticae were similar between neighboring crops, even if the acaricide had been registered for use only in almonds (cyhexatin) or cotton (dicofol). Thus, longterm acaricide selection and movement of spider mites between the two crops resulted in similar proportions of resistant individuals. In these study sites, large-scale dispersal ofT. pacificus from almonds rarely directly affected acaricide efficacy in cotton, because resistance frequencies were similar for spider mites from the two crops and because acaricide applications were usually made in cotton after dispersal from almonds was completed. In two cotton sites, field selection with dicofol was reversed by subsequent immigration of spider mites from neighboring field crops.     

Neoseiulus fallacis: dispersal and biological control of Tetranychus urticae following minimal inoculations into a strawberry field

In three separate tests, 100 adult female Neoseiulus fallacis (Garman) (plus immatures) were released at five point locations across 1.6-m rows of strawberries to control twospotted spider mites, Tetranychus urticae Koch. Beginning in April, during 6–12 weeks, predators controlled pests locally and dispersed downwind by 20–30 m. Up to 100 m² around each release point was colonized, and the entire 2.5 ha field was covered by predators by September. Distances dispersed by N. fallacis were similar within and across rows, suggesting that dispersal was primarily by aerial rather than by ambulatory means. Factors that affected dispersal were temperature, wind direction, density of spider mites, and mowing and flailing of foliage. An exponential model of dispersal was fitted to the data. On average, the area dispersed by N. fallacis doubled every 70 degree-days. From these results, a strategy of minimum release is suggested. To establish N. fallacis over a field in a single season, ca. 100 adult females per 1–2 m of row can be released before 1 July, after T. urticae have achieved 2–5 female adults per leaf. Releases should be 50 m apart and to the upwind side of the field. Selective sprays may be needed to suppress spider mites until predators gain control and disperse over the field.     

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.     

Effects of mite age, mite density, and host quality on aerial dispersal behavior in the twospotted spider mite

Mite age, population density, and host leaf quality affect various life history traits in spider mites. We investigated the effects of these factors on the aerial dispersal behavior of adult female twospotted spider mites,Tetranychus urticae Koch (Acari: Tetranychidae). The proportion of adult females exhibiting the dispersal behavior dropped significantly with age following adult emergence, particularly in the first 3 days. Sixty to eighty percent of female mites 2-days old or younger displayed the behavior under test conditions, whereas less than 20% of female mites older than 3-days-old showed the behavior. Younger adult females also exhibited shorter latency for the behavior, although this trend was not as clear. Leaf quality experienced during deutonymph development had no effect on the behavior adults subsequently displayed. On the other hand, adult females that fed on poor quality leaves after emergence were twice as likely to display the behavior (90% vs. 45%), and with shorter latency (37 vs. 77 min), than those that fed on high quality leaves. When newly emerged adult females encountered high mite density and dry leaves, the incidence of the behavior increased (69% vs. 47%) and latency decreased (69 vs. 93 min) compared to mites that encountered low density on well watered leaves. Our results suggest that both starvation and desiccation of adult females may enhance their dispersal behavior.     

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.     

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.     

Dispersal of diapausing Tetranychus urticae and T. kanzawai

The spider mites Tetranychus urticae Koch and Tetranychus kanzawai Kishida (Acari: Tetranychidae) overwinter mostly as mated adult diapausing females. Their overwintering survival depends in part on their dispersal towards suitable habitats. We investigated the dispersal behaviour of diapausing females of T. urticae and T. kanzawai with respect to factors known to affect the dispersal of non‐diapausing mites: light, population density, gravity, and humidity. In general, diapausing females of T. urticae showed a stronger tendency to disperse than did those of T. kanzawai under all test conditions. High population density promoted the dispersal of diapausing T. urticae, but not of T. kanzawai. Dispersal of diapausing females of both species was not significantly affected by gravity, humidity, or whether feeding damage was caused by conspecifics or heterospecifics. On plants, more T. urticae than T. kanzawai moved downward. We propose that dispersal after the onset of diapause may be an important life‐history strategy in T. urticae, but not in T. kanzawai.     

Effects of acaricides,pyrethroids and predator distributions on populations of <Emphasis Type="Italic">Tetranychus urticae</Emphasis> in apple orchards

We sampled mites in three apple orchards in Nova Scotia, Canada, that had been inoculated with pyrethroid-resistant Typhlodromus pyri and had a history of Tetranychus urticae outbreaks. The objective of this study was to monitor populations of T. urticae and phytoseiid predators on the ground and in trees and to track dispersal between the two habitats. Pesticides were the chief cause of differences in mite dynamics between orchards. In two orchards, application of favourably selective acaricides (abamectin, clofentezine) in 2002, coupled with predation by T. pyri in trees and Neoseiulus fallacis in ground cover, decreased high T. urticae counts and suppressed Panonychus ulmi. By 2003 phytoseiids kept the tetranychids at low levels. In a third orchard, application of pyrethroids (cypermethrin, lambda-cyhalothrin), plus an unfavourably selective acaricide (pyridaben) in 2003, suppressed phytoseiids, allowing exponential increases of T. urticae in the ground cover and in tree canopies. By 2004 however, increasing numbers of T. pyri and application of clofentezine strongly reduced densities of T. urticae in tree canopies despite high numbers crawling up from the ground cover. Another influence on T. urticae dynamics was the distribution of the phytoseiids, T. pyri and N. fallacis. When harsh pesticides were avoided, T. pyri were numerous in tree canopies. Conversely, only a few N. fallacis were found there, even when they were present in the ground cover and on tree trunks. Low numbers were sometimes due to pyrethroid applications or to scarcity of prey. Another factor was likely the abundance of T. pyri, which not only competes with N. fallacis, but also feeds on its larvae and nymphs. The scarcity of a specialist predator of spider mites in trees means that control of T. urticae largely depends on T. pyri, a generalist predator that is not particularly effective in regulating T. urticae. The Canadian Crown's right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

Phytoseiid dispersal at plant to regional levels: a review with emphasis on management of Neoseiulus fallacis in diverse agroecosystems

Dispersal behaviors of phytoseiid and tetranychid mites are key factors in understanding predator-prey dynamics and biological control of pest mites at different spatial levels in agricultural and natural ecosystems. In this review, ambulatory and aerial dispersal of both mite groups are discussed at spatial levels of leaf, plant, crop and region. Emphasis is on dispersal of phytoseiids, and specifically, the specialist-predator, Neoseiulusfallacis (Garman), and two-spotted spider mite prey, Tetranychus urticae (Koch). Dispersal aspects that are discussed are ambulation on a leaf; plant or in a prey patch; aerial dispersal between plants; behavior and aerodynamics of aerial take-off; modeling vs. monitoring of dispersal distance; fates of dispersing mites that land on soil substrates; plants as take-off platforms and landing targets for dispersers; and regional dispersal patterns and integrated mite management.     

Effects of mating status and age on dispersal behavior in the twospotted spider mite, Tetranychus urticae in response to fenvalerate-treated leaf surfaces

Effects of variation in age and mating status on dispersal of the twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), from fenvalerate-treated leaf surfaces were investigated. Unmated and mated female mites, 1 to 6 days in age, were scored for walkoff and spindown dispersal response on bean leaves treated with a sub-lethal, residual dose of fenvalerate. Variation in mite age did not have a pronounced effect on dispersal response. Significant differences in dispersal response due to mating type were observed, yet the mean differences in dispersal response between unmated and mated females was generally small. Our data suggest that the current practice of using mated female mites of unknown age when testing for among-population differences in dispersal response, in most cases, provides an adequate estimate of population response when differences among treatments in dispersal response are large. However, when testing for within-population differences in dispersal response, small treatment effects may be more easily measured using a single age class of mites, because this could decrease within-treatment variance.     

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.     

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.     

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.     

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.     

Influence of diet on life table parameters of <Emphasis Type="Italic">Iphiseius degenerans</Emphasis> (Acari: Phytoseiidae)

Studies on the reproduction, longevity and life table parameters of Iphiseius degenerans (Berlese) were carried out under laboratory conditions of 25 ± 1 °C, 75 ± 5% RH and 16L:8D h. As food sources for the predatory mite, Ricinus communis L. pollen, all stages of the spider mite Tetranychus urticae Koch, Frankliniella occidentalis (Pergande) larvae, and Ephestia kuehniella Zeller eggs were selected. All diets were accepted as food by the adult mites. Female longevity ranged from 29.5 to 42.4 days, the highest value was recorded on a diet of Ephestia eggs. The highest percentage of females escaping the experimental arena was observed on the diet consisting of thrips larvae. The highest oviposition rate (1.9 eggs/female.day) was recorded when the predator was fed on spider mites on an artificial substrate. For other diets, oviposition rates ranged from 1.0 to 1.3 eggs/female.day. The intrinsic rate of natural increase (r _m) of I. degenerans varied between 0.015 and 0.142 females/female.day. The diet consisting of castor bean pollen resulted in the highest population growth whereas the diet on spider mites brushed off onto a bean leaf arena resulted in the slowest population growth. This can be explained by the inability of the predator to cope with the webbing of T. urticae, and the high escape rate of the progeny when reared on spider mites. The percentage of females in the offspring ranged from 40 to 73%.This revised version was published online in May 2005 with a corrected cover date.     

Evidence of a high level of gene flow among apple trees in <Emphasis Type="Italic">Tetranychus urticae</Emphasis>

The dispersal mechanism of the two-spotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae) could affect predator–prey population dynamics and the spread of acaricide resistance. To investigate the propensity for spider mite migration in the field, the genetic structure of spider mite populations was studied in two apple orchards using five microsatellite markers. Adult female mites were collected from trees separated by approximately 10–24 m along a line covering a distance of about 100 m. The genetic data suggested that a high population density increased the migration rate among the breeding colonies within a single tree. Spatial autocorrelation analysis suggested a positive genetic structure in the first distance class within the two orchards, which might have been caused by crawling or short-distance aerial dispersal. Meanwhile, mites may also have a large-scale migration system that could cause a high level of gene flow and constrained isolation-by-distance or genetic clines within the approximately 100-m range of the study sites. Therefore, mites might aerially disperse over long distances on a scale of <100 m while also taking shorter trips among nearby trees within a distance of 10–24 m in the apple orchards.     

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.     

Spider-Mite Problems and Control in Taiwan

Problems with spider mites first appeared in Taiwan in 1958, eight years after the importation of synthetic pesticides, and the mites evolved into major pests on many crops during the 1980s. Of the 74 spider mite species recorded from Taiwan 10 are major pests, with Tetranychus kanzawai most important, followed by T. urticae, Panonychus citri, T. cinnabarinus, T. truncatus and Oligonychus litchii. Most crops suffer from more than one species. Spider mites reproduce year-round in Taiwan. Diapause occurs only in high-elevation areas. Precipitation is the most important abiotic factor restricting spider-mite populations. Control is usually accomplished by applying chemicals. Fifty acaricides are currently registered for the control of spider mites. Acaricide resistance is a serious problem, with regional variation in resistance levels. Several phytoseiid mites and a chrysopid predator have been studied for control of spider mites with good effect. Efforts to market these predators should be intensified so that biological control can be a real choice for farmers.