Aerial dispersal of phytoseiid mites (Acari: Phytoseiidae): estimating falling speed and dispersal distance of adult females

Aerial dispersal is important to immigration and redistribution of phytoseiid mites that often can provide biological control of spider mite pests. Falling speed of a mite and wind largely determine dispersal distance of such a passively blown organism. A diffusion model of wind-blown phytoseiids could provide insight into their dispersal. To this end, we measured body weights and falling speeds of adult females of 13 phytoseiid and one tetranychid mite species. These data were then incorporated into seed dispersal models (Greene and Johnson, Okubo and Levin) and results were compared to mite dispersal distances in wind tunnel, greenhouse and field. Weights of phytoseiid species ranged from 5.25 to 2l.7 μg; starved mites weighed less than fed mites. Geometric diameters ( d _g ) of idiosomas were correlated to weights. Falling speeds for phytoseiids were 0.39–0.73 m/s, and less than for T. urticae (0.79 m/s) in still air. In some species, active mites had slower falling speeds than inactive (anesthetized) mites indicating that behavior may influence falling. Starved mites had significantly slower falling speeds than fed mites and dispersed farther. Equation-based estimates of falling speed were close to measured ones (2–8% deviation) for some species. There were significant relationships between falling speed and body weight and morphological traits. Greene and Johnson's seed dispersal model provided better fits to dispersal of mites in the wind tunnel, greenhouse and field studies than Okubo and Levin's model. Limits of models in describing mite dispersal distance and applications to IPM are discussed.     

Relationship between dispersal and fecundity ofTetranychus urticae Koch (Acari: Tetranychidae) exposed to flucythrinate

The effects of the synthetic pyrethroid flucythrinate on dispersal and fecundity of the two-spotted spider mite,Tetranychus urticae Koch, was investigated using a vial exposure technique that allowed treatment of mites without incidental treatment of host plant and allowed determination of the actual dose received by the mites. Flycythrinate elicited both immediate and delayed dispersal. Immediate dispersal within 120 min posttreatment was more sensitive than delayed dispersal and fecundity at 24, 48, and 72 h posttreatment. The lowest flucythrinate concentrations elicited immediate dispersal without subsequent inhibition of fecundity, providing support for the hypothesis that pyrethroid-induced mite population outbreaks can result from dispersal of healthy mites to areas of low competition. Delayed dispersal, however, was observed only in mites that had reduced fecundity. No significant increases in egg production were observed; however, mites exposed to the lowest concentrations of flucythrinate showed a trend toward increased fecundity at 48 and 72 h posttreatment. Since the mites were treated rather than the host plant, it appeared that the dispersal associated with flucythrinate was manifested primarily through a general stimulation of locomotor activity rather than directional locomotor response.Contribution from the Missouri Agricultural Experiment Station, Columbia, MO, Journal Series No. 10037.     

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.     

Behaviour of coconut mites preceding take-off to passive aerial dispersal

For more than three decades the coconut mite Aceria guerreronis Keifer is one of the most important pests of coconut palms and has recently spread to many coconut production areas worldwide. Colonization of coconut palms is thought to arise from mites dispersing aerially after take-off from other plants within the same plantation or other plantations. The underlying dispersal behaviour of the mite at take-off, in the airborne state and after landing is largely unknown and this is essential to understand how they spread from tree to tree. In this article we studied whether take-off to aerial dispersal of coconut mites is preceded by characteristic behaviour, whether there is a correlation between the body position preceding aerial dispersal and the direction of the wind, and whether the substrate (outer surface of coconut bracts or epidermis) and the wind speed matter to the decision to take-off. We found that take-off can sometimes be preceded by a raised body stance, but more frequently take-off occurs while the mite is walking or resting on its substrate. Coconut mites that become airborne assumed a body stance that had no relation to the wind direction. Take-off was suppressed on a substrate providing food to coconut mites, but occurred significantly more frequently on the outer surface of coconut bracts than on the surface of the fruit. For both substrates, take-off frequency increased with wind speed. We conclude that coconut mites have at least some degree of control over take-off for aerial dispersal and that there is as yet no reason to infer that a raised body stance is necessary to become airborne.     

Aerial dispersal of European red mites (Acari: Tetranychidae) in commercial apple orchards

Aerial dispersal of European red mite, Panonychus ulmi (Koch), in commercial apple orchards was estimated by trapping windborne mites. Studies were conducted at four orchards in eastern New York during 1989 and 1990 and at three orchards in western New York during 1989. In each orchard mites were trapped in three locations; the interior of the orchard, at the border of the orchard and in a field or woodlot beyond the orchard. Large numbers of mites were captured, even when the numbers of mites on apple foliage were well below levels where mite injury to leaves was visible (less than five per leaf). The log numbers of mites trapped were linearly related to the log density of mites on leaves and this relationship was consistent for each year and region the study was conducted. The trap captures among the three locations in and outside an orchard were highly correlated. The implications these findings may have on metapopulation dynamics and resistance to acaricide dynamics are discussed.     

Wheat curl mite (Acari: Eriophyidae) dispersal and its relationship with kernel red streaking in maize

Wheat curl mites, Aceria tosichella Keifer, dispersing from wheat (Triticum spp.) to nearby corn (Zea mays L.) fields play a role in the development of kernel red streaking in corn. These studies were undertaken to verify the relationship of wheat curl mite to kernel red streaking, to determine whether wheat is the main source of curl mites dispersing into corn and to determine whether planting corn in temporal or spatial isolation of wheat is a valid management strategy. These studies were conducted on farm fields using sticky traps to monitor mites, followed by sampling mature grain for kernel streaking in southwestern Ontario from 1999 to 2002. The dominant source mites were winter wheat. Mite dispersal occurred during the first 3 wk of winter wheat maturation after the wheat had reached Zadoks stage 87. Mite dispersal corresponded to prevailing winds in the area with the lowest number of mites and the lowest severity of kernel red streaking occurring 60 m from wheat fields planted to the north, south, and east of cornfields and 90 m from wheat fields planted to the west of cornfields. The severity of kernel red streaking was positively correlated with the density of wheat curl mites in corn; however, the correlation was weak and kernel red streaking was still high in many cornfields when few or no mites were present. These findings suggest that wheat curl mite migration into corn is not entirely predictive of the incidence and severity of kernel red streaking.     

Spatio-temporal population genetic structure of the parasitic mite Spinturnix bechsteini is shaped by its own demography and the social system of its bat host

Information about the population genetic structures of parasites is important for an understanding of parasite transmission pathways and ultimately the co-evolution with their hosts. If parasites cannot disperse independently of their hosts, a parasite's population structure will depend upon the host's spatial distribution. Geographical barriers affecting host dispersal can therefore lead to structured parasite populations. However, how the host's social system affects the genetic structure of parasite populations is largely unknown. We used mitochondrial DNA (mtDNA) to describe the spatio-temporal population structure of a contact-transmitted parasitic wing mite ( Spinturnix bechsteini ) and compared it to that of its social host, the Bechstein's bat ( Myotis bechsteinii ). We observed no genetic differentiation between mites living on different bats within a colony. This suggests that mites can move freely among bats of the same colony. As expected in case of restricted inter-colony dispersal, we observed a strong genetic differentiation of mites among demographically isolated bat colonies. In contrast, we found a strong genetic turnover between years when we investigated the temporal variation of mite haplotypes within colonies. This can be explained with mite dispersal occuring between colonies and bottlenecks of mite populations within colonies. The observed absence of isolation by distance could be the result from genetic drift and/or from mites dispersing even between remote bat colonies, whose members may meet at mating sites in autumn or in hibernacula in winter. Our data show that the population structure of this parasitic wing mite is influenced by its own demography and the peculiar social system of its bat host.     

Species status and population genetic structure of grapevine eriophyoid mites

Bud mite, blister mite (Colomerus vitis Pagenstecher), and rust mite (Calepitrimerus vitis Nalepa) (Acari: Eriophyoidae) are recognized grapevine pests. Much of the biology and ecology of these pests is poorly understood. We used two types of molecular markers to gain further insight into the breeding biology and population structure of these mites, using individuals collected from sites around south‐eastern Australia. Patterns of genetic variation observed using PCR‐RFLP of ITS 1 (Internal Transcribed Spacer 1) confirmed the separate species status of the rust mite, and resolved the species status of bud and blister mites, revealing two closely related but distinct species. Microsatellite markers revealed extensive genetic differentiation between bud mite populations and blister mite populations even at micro‐geographical levels, suggesting low movement in these species. The findings indicate that separate control strategies are needed against bud and blister mites, and that localized control strategies are likely to be effective given their limited dispersal.     

Dispersal of the broad mite, Polyphagotarsonemus latus (Acari: Tarsonemidae) on Bemisia argentifolii (Homoptera: Aleyrodidae)

The broad mite Polyphagotarsonemus latus (Banks) and silverleaf whitefly Bemisia argentifolii Bellows & Perring (=B strain of Bemisia tabaci (Gennadius)) have many common host plants. It was found that broad mites can attach themselves to B. argentifolii adults and use them as a carrier for their dispersal. In a cage experiment, we observed that more than 80% of B. argentifolii adults had more than one broad mite attached within 4 h after B. argentifolii landed on broad mite-infested plants. Overall, 97.5% of the broad mites examined were attached to the legs, mostly on the tibiae and tarsi of B. argentifolii adults, and 99.5% of the broad mites attached to B. argentifolii were adult females. The successful dispersal of broad mite via B. argentifolii was also demonstrated with a cage experiment.     

Of mites and movement: the effects of plant connectedness and temperature on movement of Phytoseiulus persimilis

Simulation modelling studies on the biological control of Tetranychus urticae Koch in ornamental crops suggest that the dispersal of the predatory mite Phytoseiulus persimilis Athias-Henriot in the absence of food is important in determining its ability to locate sparsely distributed patches of prey (Skirvin et al., 2002). Experimental work to examine factors influencing dispersal of P. persimilis has shown that ground substrate affects the movement of the predator, and that the greater the number of connections between adjacent plants the greater the number of mites moving. In addition, P. persimilis are able to move across as many as 10 plant–plant connections within 24 h, although the majority of predators tracked moved less than this. Temperature has a significant impact on dispersal of P. persimilis, with more mites leaving release points as temperature increases up to 25 °C, but decreasing above this temperature. This work highlights the importance of understanding how the plant canopy and temperature influence the dispersal of predatory mites. The importance of these results for biological control in ornamental crops is discussed.     

Biological control of sciarid and phorid pests of mushroom with predatory mites from the genus Hypoaspis(Acari: Hypoaspidae) and the entomopathogenic nematode Steinernema feltiae

In small-scale experiments, the predatory mites, Hypoaspis aculeifer (Canestrini) and H. miles Berlese, applied at 700 mites m(-2), and the entomopathogenic nematode, Steinernema feltiae (Filipjev) applied at 3 x 10(-6) nematodes m(-2) controlled sciarids and phorids in mushroom compost and casing substrates. For both mite species, earliest application to the growing substrate following sciarid infestation reduced sciarid emergence. In contrast, later application of each biological control agent provided more effective control of phorid emergence. The behaviour of adult mites suggested that H. aculeifer were more positively geotactic than H. miles although both species could penetrate compost and casing substrates to a depth of 2-12 cm. A majority of S. feltiae nematodes resided at a depth of 2-4 cm in both substrate types. Independent application of H. aculeifer provided more comprehensive control of sciarids and phorids than the other biological agents studied, owing to its better dispersal within compost and casing, and ability to attack larvae of differing ages.     

Ambulatory dispersal behavior ofNeoseiulus fallacis (Acarina: Phytoseiidae) in relation to prey density and temperature

Ambulatory dispersal behavior ofNeoseiulus fallacis (Garman) was studied in the laboratory to evaluate within-plant movement in relation to temperature and prey density. Adult femaleN. fallacis were confined in 2.5-cm-diameter arenas on the abaxial surface of excised corn leaves. Four temperatures (23, 28, 33, and 39° C) and prey densities ranging from 0 to 55 spider mite eggs per cm² were used. The walking paths of these mites were traced, digitized and used to calculate turning angles, walking speeds and turning rates. A computer simulation of walking behavior used this information to model mite ambulatory behavior and predict dispersal rates.Neoseiulus fallacis behavior while on whole corn leaves was quantified to verify the results of the simulation. The results showed thatN. fallacis will follow a leaf or arena edge (edge-walking) at all temperatures and prey densities. In addition, this behavior was used to the exclusion of the other types of behavior such as resting, and random-walk type search when prey egg density was less than 4 eggs per cm². The exclusion of edge-walking behavior from the model caused the model to underestimate substantially the dispersal rates leaves. These data suggest that there are at least two recognizable types of ambulatory search used byN. fallacis—the random-walk type, which is used when prey density is high (searching within prey patches), and the edge-walking behavior, which is used when prey density is low. This behavior allows the mite to travel rapidly from leaf to leaf in search of new prey patches.     

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.     

Population density and phenology of Tetranychus urticae (Acari: Tetranychidae) in hop is linked to the timing of sulfur applications

The twospotted spider mite, Tetranychus urticae Koch, is a worldwide pest of numerous agronomic and horticultural plants. Sulfur fungicides are known to induce outbreaks of this pest on several crops, although mechanisms associated with sulfur-induced mite outbreaks are largely unknown. Studies were conducted during 2007-2009 in Oregon and Washington hop yards to evaluate the effect of timing of sulfur applications on T. urticae and key predators. In both regions, applications of sulfur made relatively late in the growing season (mid-June to mid-July) were associated with the greatest exacerbation of spider mite outbreaks, particularly in the upper canopy of the crop. The severity of mite outbreaks was closely associated with sulfur applications made during a relatively narrow time period coincident with the early exponential phase of spider mite increase and rapid host growth. A nonlinear model relating mean cumulative mite days during the time of sulfur sprays to the percent increase in total cumulative mite days (standardized to a nontreated plot) explained 58% of the variability observed in increased spider mite severity related to sulfur spray timing. Spatial patterns of spider mites in the Oregon plots indicated similar dispersal of motile stages of spider mites among leaves treated with sulfur versus nontreated leaves; however, in two of three years, eggs were less aggregated on leaves of sulfur-treated plants, pointing to enhanced dispersal. Apart from one experiment in Washington, relatively few predatory mites were observed during the course of these studies, and sulfur-induced mite outbreaks generally occurred irrespective of predatory mite abundance. Collectively, these studies indicate sulfur induces mite outbreaks through direct or indirect effects on T. urticae, mostly independent of predatory mite abundance or toxicity to these predators. Avoidance of exacerbation of spider mite outbreaks by sulfur sprays was achieved by carefully timing applications to periods of low spider mite abundance and slower host development, which is generally early to mid-spring for hop.     

Recapture of male and female dragonflies in relation to parasitism by mites, time of season, wing length and wing cell symmetry

For aquatic mites parasitic on dragonflies, completion of their life cycle depends on their being returned to appropriate water bodies by their hosts, after completion of engorgement. We examined whether differences among hosts in timing of emergence or phenotypic attributes might affect their probability of return to an emergence pond, and hence success of mites. Parasitized males and females of the dragonfly Sympetrum obtrusum (Hagen) did not differ in overall recapture rates. Females that had wing cell symmetry and emerged early were more likely to be recaptured than females that emerged later or had wing cell asymmetry, but there were no consistent relations between these variables and parasitism by mites. No such relations between wing cell asymmetry, emergence date, and recapture likelihood were found for males. Using randomization tests, we found that mean intensities of Arrenurus planus (Marshall) mites at host emergence were the same for recaptured females and females not recaptured; however, males that were recaptured had lower mean intensities of mites at emergence than males not recaptured. Further, mature females carried more mites than mature males, and the latter had fewer mites than newlyemerged males not recaptured. Biases in detachment of engorging mites do not explain the differences in parasitism between mature males and females, nor the differences in mite numbers between mature males and newly emerged males that were not recaptured. Rather, heavily parasitized males appear to disperse or die and are not recaptured, which should have implications for dispersal of mites and fitness of male hosts.     

An acarine predator-prey metapopulation system inhabiting greenhouse cucumbers

The two-spotted spider mite ( Tetranychus urticae ) is a serious pest on greenhouse cucumbers, but can be controlled by the phytoseiid predator Phytoseiulus persimilis. The two mite species exhibit considerable fluctuations in overall population densities but within acceptable limits. The system appears to be persistent at a regional (greenhouse) scale in spite of frequent local extinctions (e.g. at individual plants). Experimental evidence indicates that the mites form a metapopulation system characterized by 'shifting mosaic' dynamics. A stochastic simulation model is used to analyse the role played by dispersal in the dynamics and persistence of the system. It shows that demographic stochasticity generates sufficient endogenous 'noise' to counteract the synchronizing effect of density-dependent dispersal, provided dispersal rates are not too high and the system is not too small. Low dispersal rates, on the other hand, increase the risk of local outbreaks of spider mites that may cause destruction of plants.     

Simulation of the interactions of predatoryTyphlodromus mites with the European red mite,Panonychus ulmi (Koch)

Interactions of Typhlodromus mites and their prey, the European red mite, in orchard settings were examined through computer simulation. In particular, the consequences of very slow dispersal by the predators, compared with the weather-dependent higher rates of prey dispersal, were elucidated. In simulations of the interactions of the predator and its prey, both dispersal and temperature strongly affected the available supply of juvenile prey, and thus could determine whether the predators on an individual tree survived or perished.     

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.     

Dispersal strategies of Aceria guerreronis (Acari: Eriophyidae), a coconut pest

The dispersal of plant-feeding mites can occur involuntarily, through transportation of infested plant parts, or voluntarily, by walking to new plant parts or to suitable spots where biotic (phoresis) or abiotic (wind, agricultural tools, etc.) factors carry them over long distances. Elucidating the dispersal mechanisms of the coconut mite, Aceria guerreronis Keifer, is important for understanding the process of colonization of new fruits of a same or different plants, essential for the improvement of control strategies of this serious coconut pest. Thus, the objective of this work was to investigate the voluntary dispersal mechanisms of this mite. The hypothesis that the coconut mite disperses by walking, phoresis or wind were tested. The coconut mite was shown to be able to walk short distances between fruits of the same bunch or between bunches of the same plant. Phoresis on insects of the orders Hymenoptera (Apidae), Coleoptera (Curculionidae) and Lepidoptera (Phycitidae) was evaluated in the laboratory and in the field. Although in the laboratory mites were shown to be able to climb onto honeybees, field investigations failed to show these insects as important carriers of the pest, corroborating findings of previous works; however, both laboratory and field investigations suggested the curculionid Parisoschoenus obesulus Casey to be able to transport the coconut mite between plants. Similarly, laboratory and field investigations suggested wind to be important in the dispersal of the coconut mite between plants.     

Correlation between Olfactory Responses, Dispersal Tendencies, and Life-history Traits of the Predatory Mite Neoseiulus Womersleyi (Acari: Phytoseiidae) of Eight Local Populations

To investigate the relationship between foraging behavior and life-history traits of the predatory mite Neoseiulus womersleyi, the olfactory responses, dispersal ratios from a prey patch, predation rates, fecundity, and developmental times in eight local populations of N. womersleyi were investigated. Significant differences among local populations were found in all these traits except fecundity. None of the life-history traits correlated with foraging behavior. A significant positive correlation was found only between the olfactory response and the dispersal ratio. These results suggested that predatory mites with low olfactory responses would stay in a prey patch longer than predatory mites with high olfactory responses.