Kampimodromus aberrans (Acari: Phytoseiidae) from the USA: morphological and molecular assessment of its density.

Morphological measurements and a mitochondrial molecular marker (COI) were used to identity specimens reported as Kampimodromus aberrans on hazelnut in the USA. Several species and populations of this genus were studied to assist with identification. Both data types showed that specimens from the USA differed from K. aberrans from other regions. USA specimens seem to belong to the same species as Kampimodromus specimens from France on hazelnut. These mites were morphologically similar to Kampimodromus coryli and K. corylosus, which according to the original descriptions, are distinguished by the presence or absence of a tooth on the movable digit of the chelicera, with K. coryli having one tooth and K. corylosus none. As chelicerae of Kampimodromus from hazelnut in the USA and France are toothless, they are assigned to the species K. corylosus. Studies showed that morphological characters traditionally used to identify Kampimodromus species, such as setal length, are of less value than other characters that are difficult to observe, such as the numbers of solenostomes and the presence of teeth on the movable digit of the chelicerae. Some synonyms are discussed.

Evaluation of prey-stage preference as an indicator of life-style type in phytoseiid mites.

Discriminant analysis (DA) models were developed and applied to examine the use of prey-stage preference (Tetranychus urticae Koch egg versus larval prey) in the classification of phytoseiid mites into life-style types. Prey-stage preferences and developmental times when preying on T. urticae, and relative ovipositional rates on six food categories were determined for four phytoseiid species occurring on apple in central and eastern Oregon, USA: Galendromus flumenis (Chant), Galendromus occidentalis (Nesbitt), Metaseiulus citri (Garman and McGregor) and Typhlodromus caudiglans Schuster. In terms of all three aspects studied, the phytoseiid species showed a consistent polarization of G. occidentalis < or = G. flumenis < or = T. caudiglans < M. citri. Specifically, G. occidentalis ('The Dalles' strain) had a significant preference for eggs, G. flumenis had no preference, and T. caudiglans and M. citri had significant preferences for larvae; G. occidentalis had the shortest developmental time, followed by G. flumenis and T. caudiglans, while M. citri had the longest developmental time; and diet breadth was most narrow for G. occidentalis and progressively broader from G. flumenis, T. caudiglans through M. citri, which was able to sustain oviposition on the broadest range of prey and pollens. Species were classified somewhat differently depending on which traits were considered in a given DA. Prey-stage preference was not included as an indicator in the parsimonious DA model when all species and all traits were considered, but in general this trait performed well as an indicator alone (single-trait DA) and somewhat improved the classifications of multitrait discriminant analyses.

Phytoseiid mites on unsprayed apple trees in Oregon, and other western states (USA): distributions, life-style types and relevance to commercial orchards.

In unsprayed apple trees in eastern Oregon, Galendromus flumenis (Chant), Galendromus occidentalis (Nesbitt), Typhlodromus caudiglans Schuster and Metaseiulus citri (Garman and McGregor) were common phytoseiid mites; common plant-feeding mites were the eriophyid, Aculus schlechtendali Nalepa, the brown mite, Bryobia rubrioculus (Scheuten) and Eotetranychus spp.; apple rust mites seemed to be the primary prey for phytoseiids; the spider mites, Tetranychus urticae Koch and Panonychus ulmi (Koch) were scarce except for a few local outbreaks; the stigmaeid Zetzellia mali (Ewing) was at 10% of sites and its densities were inversely related to phytoseiid densities; phytoseiids were absent at some sites, particularly at high elevations where winters are severe. In seven Oregon ecoregions, G. flumenis was often at lower elevations in valleys with moderate winters; T. caudiglans was often at higher elevations; G. occidentalis was often at intermediate elevations, in young trees, and near where pesticides were used; it dominated in unsprayed trees only in almost treeless, sage-covered areas; M. citri was usually in older apple trees near agriculture. In mixed phytoseiid populations, M. citri, a generalist, and G. occidentalis, a specialist, occurred more often than expected; G. occidentalis was mostly found with T. caudiglans, a generalist; G. flumenis, a generalist, occurred less with others, possibly because it competes with both specialists and generalists. Analyses of species' distributions with multiple regression and genetic models gave explanatory r2s of 0.019-0.318. Of 29 variables, altitude of site, intensity of agricultural management, tree age, plant types, and Z. mali levels helped explain phytoseiid species presence. In the western USA, G. flumenis dominated in middle-southern latitudes; T. caudiglans dominated in the north near the Canadian border; G. occidentalis dominated in middle latitudes in parts of Washington, Idaho, Montana, and Wyoming; M. citri was at a few sites in these four states. Distributional and independent variable data were used to predict species presence at sites in Oregon with a 70% success rate. We discuss phytoseiid life-style types, community dynamics, presence in organic/conventional orchards, and trends as more selective IPM methods are used in apple orchards.

Low-density releases of Neoseiulus fallacis provide for rapid dispersal and control of Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae) on apple seedlings.

Releases of Neoseiulus fallacis (Garman) at 1500--6000 per ha when prey were at 0.1-0.3 per leaf provided seasonal control of Tetranychus urticae Koch (all stages) at 1-2 per leaf in an apple seedling rootstock nursery. Predaceous mites (all stages) increased to 0.3-0.4 per leaf after releases and predator prey ratios of < or = 1:3-7 provided pest regulation thereafter. Such low-density releases were thought to be effective because multiple dispersal bouts allowed predators to locate widely distributed spider mites (on 2-6% of leaves). A random-diffusion model simulating predator dispersal (incorporating wind speed and direction parameters) adequately explained movement and pest control patterns. An upright, dense, uniform planting of apple seedlings was an effective producer and recipient for dispersing predators and these attributes seemed to explain why biological control was so effective. Low-density releases of N. fallacis for control of T. urticae are predicted to be less effective on other crops with less prominent profiles and soil coverage.

Biological control of spider mites on grape by phytoseiid mites (Acari: Tetranychidae, Phytoseiidae): emphasis on regional aspects.

Leaf samples were taken from 34 (1998) and 10 (1999) vineyards in five valleys in western Oregon to assess spider mite pests and biological control by predaceous phytoseiid mites. A leaf at a coordinate of every 10 m of border, 5 m into a vineyard, was taken to minimize edge effects; 20 leaves were taken at regular intervals from vineyard centers. Variables recorded at each site included grape variety and plant age, chemicals used, and vegetation next to vineyards. Sites were rated as occurring in agricultural versus riparian settings based on surrounding vegetation types. Multiple linear regressions and a computer genetic algorithm with an information content criterion were used to assess variables that may explain mite abundances. Typhlodromus pyri Scheuten was the dominant phytoseiid mite species and Tetranychus urticae Koch the dominant tetranychid mite species. High levels of T. urticae occurred when phytoseiid levels were low, and low levels of T. urticae were present when phytoseiid levels were high to moderate. T. urticae densities were higher in vineyards surrounded by agriculture, but phytoseiid levels did not differ between agricultural and riparian sites. Phytoseiids had higher densities on vineyard edges; T. urticae densities were higher in centers. Biological control success of pest mites was rated excellent in 11 of 44 vineyards, good in 27, and poor in only six sites. Predaceous mites appeared to be the principal agents regulating spider mites at low levels in sites where pesticides nontoxic to predators were used. Effects of surrounding vegetation, grape variety, growing region, and other factors on mites are discussed.

Plant-related factors influence the effectiveness of Neoseiulus fallacis (Acari: Phytoseiidae), a biological control agent of spider mites on landscape ornamental plants.

The predatory mite Neoseiulus fallacis (Garman) was evaluated as a biological control agent of herbivorous mites on outdoor-grown ornamental landscape plants. To elucidate factors that may affect predator efficiency, replicated tests were conducted on 30 ornamental plant cultivars that varied in relationship to their generalized morphology (e.g., conifers, shade trees, evergreen shrubs, deciduous shrubs, and herbaceous perennials), production method (potted or field grown), canopy density, and the prey species present on each. Plant morphological grouping and foliar density appeared to be the most influential factors in predicting successful biological control. Among plant morphological groups, N. fallacis was most effective on shrubs and herbaceous perennials and less effective on conifers and shade trees. N. fallacis was equally effective at controlling spider mites on containerized (potted) and field grown plants, and there was no difference in control of mites on plants with Tetranychus spp. versus those with Oligonychus or Schizotetranychus spp. Moderate to unsuccessful control of spider mites by N. fallacis occurred mostly on tall, vertical plants with sparse canopies. Acceptable spider mite control occurred in four large-scale releases of N. fallacis into production plantings of Abies procera, Thuja occidentalis 'Emerald', Malus rootstock, and Viburnum plicatum 'Newport'. These data suggest that N. fallacis can be an effective biological control agent of multiple spider mite species in a range of low-growing and selected higher growing ornamental plants.

Jerking in predaceous mites (Acari: Phytoseiidae) with emphasis on larvae.

Jerking, a behavior involving a pronounced and often repeated lunging of the body, was studied in larvae among seven species of phytoseiid mites, Euseius finlandicus, Galendromus occidentalis, Neoseiulus californicus, Neoseulus fallacis, Phytoseiulus macropilis, Phytoseiulus persimilis and Typhlodromus pyri, and in the nymphal stages of N. fallacis. This behavior was observed in larvae of six of the seven species and in all active immature stages of N. fallacis. Jerking was usually triggered by direct contact with a con- or heterospecific mite, although jerking occurred occasionally without direct contact in N. fallacis larvae. The larval jerking tendency (the mean fraction of contacts resulting in jerking) of a species was significantly correlated with a tendency to congregate with conspecifics. Congregated larvae often probed their immediate area and each other with the front legs, repeatedly jerking in response to contact with neighboring larvae. Species with larvae having higher jerking tendencies also jerked more repeatedly per jerking event. Among species, there was no consistent relationship between jerking tendency and larval age. Larvae of N. fallacis had a higher tendency to jerk than protonymphs and deutonymphs, and they had both the highest jerking and congregating tendencies among larvae of all species studied. An individual was more likely to jerk when approached by another mite than when it initiated contact, and when observed in N. fallacis larvae that initiated contact, jerking was sometimes aggressive. Jerking N. fallacis larvae experienced fewer and shorter periods of probing by the mouthparts and fewer attacks from cannibalistic adult females than anesthetized larvae that were unable to jerk.

Quantitative classification of life-style types in predaceous phytoseiid mites.

Classification of species into different functional groups based on biological criteria has been a difficult problem in ecology. The difficulty mainly arises because natural classification patterns are not necessarily mutually exclusive. The more group characteristics overlap, the more difficult it is to identify the membership of a species in the overlapping portions of any two groups. In this paper, we present an application of discriminant analysis by creating classification models from life history and morphological data for two specialist and two generalist life-styles type of predaceous phytoseiid mites. Two stages can be distinguished in our method: life-style group membership assignment and trait variable evaluation. We use a Bayesian framework to create a classifier system to locate or assign species within a mixture of trait distributions. The method assumes that a mixture of trait distributions can represent the multiple dimensions of biological data. The mixture is most evident near the boundaries between groups. Because of the complexity of analytical solution, an iterative method is used to estimate the unknown means, variances, and mixing proportion between groups. We also developed a criterion based on information theory to evaluate model performance with different combinations of input variables and different hypotheses. We present a working example of our proposed methods. We apply these methods to the problem of selecting key species for inoculative release and for classical introductions of biological pest control agents.

Species association among predaceous and phytophagous apple mites (Acari: Eriophyidae, Phytoseiidae, Stigmaeidae, Tetranychidae).

Predator-predator, predator-prey, and prey-prey associations among nine species of mites were studied in a plot of 100 'Red Delicious' apple (Malus pumila Miller) trees from 1990 to 1997. In 1990, seven-year-old trees were inoculated with Panonychus ulmi (Koch), Tetranychus urticae Koch (Acari: Tetranychidae) or both, and sprayed with azinphosmethyl (alone or plus endosulfan), or nothing. The species Zetzellia mali (Ewing) (Acari: Stigmaeidae), Amblyseius andersoni Chant (Acari: Phytoseiidae), Eotetranychus sp., Bryobia rubrioculus (Scheuten) (Acari: Tetranychidae), and Aculus schlechtendali Nalepa (Acari: Eriophyidae) were already present or immigrated into plots, and Galendromus occidentalis (Nesbitt) and Tvphlodromus pyri Scheuten (Acari: Phytoseiidae) were introduced. Yule's V association index was used to measure positive, neutral, or negative interspecific associations for each species pair, because of its robustness with spatially autocorrelated data. We found that pesticide and release treatments did not greatly affect the association results, but there were strong seasonal differences. Predator-predator associations were the strongest and most consistent, showing negative associations in the early and mid seasons, and neutral ones in late season. Negative associations of T pyri with other predators were the strongest, which is consistent with evidence that this mite can detect other predators on a leaf. Predatorprey seasonal associations were mixed, with some positive and others negative, with most significant associations occurring in the mid season. One prey-prey interaction was positive, again in mid season, most likely because of similar habitat preferences.

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.

Survival and plant-prey finding by Neoseiulus fallacis (Acari: Phytoseiidae) on soil substrates after aerial dispersal.

In a greenhouse and in an open field, aspects of aerial and ambulatory dispersal of the phytoseiid mite. Neoseiulus fallacis (Garman) were studied with a focus on events that would occur after aerially dispersing mites had landed on soil or associated substrates. We measured recovery of predators on lima bean plants (Phaseolus lunatus L.) that were infested with the two-spotted spider mite, Tetranychus urticae Koch. Factors thought to affect movement and colonization were distance to a receiver unit from a release (landing) point, intervening soil surfaces such as clods, gravel, fine soil and grass, and management of soil surfaces such as mulching. watering or both. In the field, the effect of distance (0.11-1.76 m) from a landing point to a receiver unit was significant, with a negative log-linear relationship. Soil surfaces such as clods and management actions such as watering with mulching allowed for more capture of predators on bean plants with prey than did other treatments. Environmental conditions greatly affected survival of N. fallacis. Predators in the field that were present on bare soil suffered high mortality (ca. 90%) at fluctuating daytime conditions of 26.4 +/- 4.8 degrees C and 56 +/- 13.4% RH. Predators only suffered 10% mortality in the greenhouse under the same setting, but under more controlled and favorable environmental conditions. Effects of environmental conditions, mode of dispersal and implications to biological control are discussed.

Cannibalism and intraguild predation among phytoseiid mites: are aggressiveness and prey preference related to diet specialization?

We tested whether specialist and generalist phytoseiid mites differ in aggressiveness and prey choice in cannibalism and intraguild predation. Specialists tested were Galendromus occidentalis, Neoseiulus longispinosus, Phytoseiulus persimilis, and P. macropilis; generalists tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, NM fallacis, and Typhlodromus pyri. Aggressiveness of cannibalistic females against larvae was not related to diet specialization except that highly aggressive species were exclusively generalists. Seldom to moderately cannibalizing species occurred in both generalist and specialist phytoseiids. In contrast to aggressiveness in cannibalism, generalists and specialists differed in aggressiveness in intraguild predation. Adult females of specialists were only slightly aggressive against heterospecific larvae, whereas adult females of all generalists except T. pyri were highly aggressive. Adult females of generalists were able to discriminate between con- and heterospecific larvae and preferentially consumed the latter when given a choice. Adult females of specialists except G. occidentalis showed no preference when given a choice between con- and heterospecific larvae. We conclude that aggressiveness in intraguild predation, species recognition and subsequent preferential consumption of heterospecifics when given a choice is common in generalist but not specialist phytoseiids. We discuss the evolutionary pathways that might have led to the difference between specialists and generalists in species discrimination.

Phytoseiid mites on peppermint and effectiveness of Neoseiulus fallacis to control Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae) in arid growing regions.

The humid-adapted species Neoseiulus fallacis (German) was the most common phytoseiid mite collected in either humid (> 100 cm annual rainfall) or arid (20-45 cm annual rainfall) mint growing regions of Washington, Oregon, Montana, Idaho, and California during 1991-1995. In experimental field plots, this predator gave excellent biological control of Tetranychus urticae Koch on mint grown under arid conditions in central Oregon when evaluated by an insecticide check method or by the caging of mites. N. fallacis is effective as a predator in arid areas probably because regular irrigation creates a humid environment in the canopy. The selective miticide propargite, when used in combination with predators, was effective at reducing high spider mite populations to below the treatment threshold faster than did N. fallacis alone.

Life-styles of Phytoseiid mites and their roles in biological control.

This review categorizes the diversity of life-styles in the Phytoseiidae, based primarily on food habits and related biological and morphological traits. The life-styles proposed are as follows: Type I, specialized predators of Tetranychus species represented by the Phytoseiulus species; Type II, selective predators of tetranychid mites (most frequently associated with species that produce dense webbing) represented by Galendromus, some Neoseiulus, and a few Typhlodromus species; Type III, generalist predators represented by some Neoseiulus species and most Typhlodromus and Amblyseius species, as well as species in all other genera about which information is available; Type IV, specialized pollen feeders/generalist predators represented by Euseius species. Consideration is given to the relative importance of each of these types in biological control and pest management programs.

Epoxide forming and degrading enzymes in the spider mite, Tetranychus urticae.

Enzymes associated with the epoxidation and epoxide hydration or glutathione conjugation pathway occurred in the herbivorous mite, Tetranychus urticae. Epoxidation of aldrin was primarily microsomal, required NADPH, was associated with a NADPH-cytochrome c reductase, and was inhibited by CO, 1-phenylimidazole and piperonyl butoxide. Trans- and cis-epoxide hydrolases resided mostly in the microsomal fraction but were localized also in the cytosol. These activities were differentially inhibited by 1,2-epoxy-3,3,3-trichloropropane, and chalcone and 4-phenylchalcone oxides. In vitro and in vivo rates of aldrin epoxidation were very similar indicating that in vitro artifacts were not impairing full enzyme measurement. This was further confirmed in experiments with enzyme stabilizers.

Detoxification enzyme differences between a herbivorous and predatory mite.

The detoxification capabilities of the predatory mite Amblyseius fallacis and its herbivorous prey Tetranychus urticae are fundamentally different. The activities of mixed-function oxidase and trans-epoxide hydrolase are higher in the prey than in the predator; those of cis-epoxide hydrolase and glutathione transferase are lower; and esterase activity is similar. Dissimilarities may be related both to differing adaptations to plant allelochemicals and to the higher respiration rate of the predator. Hydrolytic and conjugating reactions appear more important than oxidative pathways in imparting organophosphate resistance to these acarines. These resistances provide insecticide selectivity favorable to the predator and improved integrated pest control.

Integrated pest management in the U.S.: progress and promise.

In the U.S., where heavy use of insecticides has been commonplace for years, the development of proper integrated insect pest control cannot get underway unless there is a changed use pattern for such chemicals. A changed use pattern, however, cannot be accomplished without much study to establish the requirements for integrated control for each major crop situation. In this paper recent developments in a number of crop areas in the U.S. in which the necessary study has been begun are reviewed. Important phases in the development of integrated control programs include: the single tactics phase, the multitactic phase, phase, the biological monitoring phase, the modeling phase, the management and optimization phase, and the implementation phase. Several crops are discussed in relation to how far along we are in the development of practical programs of insect pest control. These are cotton, apples, alfalfa, soybeans, citrus, corn, cereal grains, tobacco and pine forests. Several of these programs have already made substantial headway, e.g., those for cotton, alfalfa, apples, tobacco, and soybeans, although the accomplishments have not been even or parellel with respect to the phases of development where progress has been good. The review of developments in these crops suggests that programs of control for individual crops and perhaps for complexes of associated crops will be developed according to specific needs of the crop, the geographic area and the pests, the technologies available and the socioeconomic and political factors of relevance. The tendency will be toward greater use of science in pest control decision-making, with extensive use of biological monitoring to establish realistic levels of threatened damage to the crop, and greater concern given to possible profit reductions and environmental disturbances of applying an insecticide, as well as the possible gain from doing so.