A comparative life history study of immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) with a review of larval feeding patterns in the family

Survival, developmental time, activity, feeding rates, and other biological aspects of immatures of Amblyseius fallacis, Amblyseius andersoni, Typhlodromus occidentalis and Typhlodromus pyri were examined in the laboratory in small arenas (2×2 cm) with different egg densities (0, 5, 10, 20 per 12 h) of the twospotted spider mite, Tetranychus urticae (Koch), at 25±1°C, ≈80% RH, and 16L: 8D photoperiod. Egg survival was high (86–100%) in all four species. Larval survival was similarly high except for T. occidentalis which all died in the absence of food. Survival rates of protonymphs and deutonymphs were also high except that up to 50% of A. andersoni died at 5 prey eggs per 1/2 day. Developmental time did not vary significantly with prey density and was similar for males and females in the oligophagous predators (A. fallacis and T. occidentalis), but was longer at lower prey densities and in females than males in the polyphagous predators (A. andersoni and T. pyri). In general, the time allocated to three active instars (=stases) decreased in the order: A. andersoni (81%), T. pyri (78%), A. fallacis (69%), and T. occidentalis (64%). The polyphagous predator species had a shorter larval stage and much longer deutonymphal stage than the oligophagous species. The proportion of time allocated to the protonymphal stage was the least variable among the four species. The interspecific differences in walking activities also appeared greater in larval and deutonymphal stages than in the protonymphal stage. The larvae of the two oligophagous predators (A. fallacis and T. occidentalis, walking activity averaging 36–49%) were more active than the two polyphagous predators (A. andersoni and T. pyri), which spent 80% or more time resting. In deutonymphs, walking activity increased in the order: T. occidentalis (14%), A. andersoni (27%), A. fallacis (43%) and T. pyri (59%). Larvae were more active during the first half of their life than the latter half. In general, most life history traits of immature A. andersoni, T. pyri, A. fallacis, and T. occidentalis are not associated with their phylogenetic relatedness or size, but with the feeding specialization of the predator species. Larval feeding patterns in Phytoseiidae are reviewed and a hypothesis about the evolution of larval feeding behavior in Phytoseiidae is proposed.     

Intraspecific competition in immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Intraspecific competition in immature Amblyseius fallacis, Amblyseius andersoni, Typhlodromus occidentalis and Typhlodromus pyri was examined in the laboratory using small cages at five different predator densities (two, four, eight, 16 and 32) in the absence and presence of prey 100 eggs of two-spotted spider mite, Tetranychus urticae (Koch), at 25 ± 1°C, 80% RH and 16L:8D photoperiod. In the absence of spider mite prey, some individuals of immature phytoseiids showed increased development and surival with increasing predator densities up to certain limits, but none survived to the adult stage, except for a single male each of A. andersoni and A. fallacis who completed development by cannibalizing on conspecifics at a density of 32 predators per cage. In the absence of spider mite prey, the mean immature survival time was independent of the initial predator density, but the variance of survival time increased with predator density. In the presence of prey, the proportion of immatures surviving to adulthood generally decreased with initial predator density and dropped sharply to almost none at the predator density of 32 for A. fallacis, eight for A. andersoni, 16 for T. occidentalis and four for T. pyri. The number of prey consumed per predator during the first day generally decreased with predator density in all four species, as prey available per predator decreased and the competition for food increased with predator density. Our data indicate that scramble competition is operating in these four species. Although cannibalism was occasionally observed, especially after the exhaustion of prey and in the generalist predators such as A. andersoni, the immatures of these phytoseiids were less influenced by the interference of conspecifics than by the increasing difficulty of finding food at high predator densities. The implications of this study for understanding phytoseiid population dynamics and their use in biological control are discussed.     

Intra- and interspecific predation on four life stage groups by the adult females of Metaseiulus occidentalis, Typhlodromus pyri, Neoseiulus fallacis and Amblyseius andersoni

Do adult females of oligophagous species such as Neoseiulus fallacis (Garman) and Metaseiulus occidentalis (Nesbitt) show less intra- and interspecific predation on phytoseiids when other foods are scarce than polyphagous species such as Amblyseius andersoni Chant and Typhlodromus pyri Scheuten? We caged single adult females of each species without food with ten of their own eggs or larvae, with ten eggs or larvae of the other species or with ten nymphs or adult females of M. occidentalis (T. pyri for M. occidentalis). We assessed the ambulatory activity, survival time, egg levels and prey loss in each test. Polyphages (in particular T. pyri) lived longer than oligophages (in particular N. fallacis) without food. The small T. pyri detected its own stages and benefited most by feeding on small active stages of other species. Amblyseius andersoni, the largest mite, fed and gained the most of any species when held with nymphs and female adults. Metaseiulus occidentalis fed on eggs of all four species to enhance survival. The large hyperactive N. fallacis gained the least from these behaviours. Each mite seemed uniquely adapted to survive conditions of scarce prey and these behaviours may explain their roles in phytoseiid mite complexes. Overall, oligophagous adult females fed less and gained less by feeding on phytoseiids than did polyphagous adult females.     

Walking,feeding and intraspecific interaction of larvae of Metaseiulus occidentalis,Typhlodromus pyri,Neoseiulus fallacis and Amblyseius andersoni held with and without eggs of Tetranychus urticae

Larvae of Metaseiulus occidentalis (Nesbitt), Typhlodromus pyri Scheuten, Neoseiulus fallacis (Garman) and Amblyseius andersoni Chant exhibited different activity levels when held on apple leaf or on tile arenas and given or not given eggs of Tetranychus urticae Koch and water (tiles only). M. occidentalis larvae held without prey exhibited high levels of walking (includes searching) during 24 hours of evaluation, whereas M. occidentalis larvae held with prey fed quickly and then became less active. Fed larvae of M. occidentalis were less active on leaves than tile. Larvae of T. pyri on leaves had a very low frequency of walking, almost never fed and quickly assumed a resting position during development. While much less active than M. occidentalis, fed and unfed T. pyri larvae walked more on tiles than leaves before resting. Larvae of N. fallacis and A. andersoni fed at low rates and were similarly active on tiles and leaves. Free water increased walking by M. occidentalis, A. andersoni and T. pyri on tile but not N. fallacis. M. occidentalis larvae interacted 5–7 times more often than larvae of the other three species. Cannibalism or scavenging was rarely seen and then only for M. occidentalis larvae. Larvae of all four mites walked, fed and interacted much more in the first 12 hours than the second 12 hours of tests, except unfed M. occidentalis. Unfed M. occidentalis larvae did not molt to protonymphs but unfed larvae of the other three species did. Unfed and fed protonymphs of all four species walked more at 4 hours after molting than larvae at 12–24 hours. Unfed and fed protonymphs of T. pyri or A. andersoni had similar walking frequencies, but unfed protonymphs of N. fallacis were more active than fed ones. Trends in larval activities are discussed relative to the life history of each species.     

Larval survival and feeding by immature Metaseiulus occidentalis,Neoseiulus fallacis,Amblyseius andersoni and Typhlodromus pyri on life stage groups of Tetranychus urticae Koch and phytoseiid larvae

When 20 newly hatched larvae either of Metaseiulus occidentalis (Nesbitt), Neoseiulus fallacis (Garman), Amblyseius andersoni Chant or Typhlodromus pyri Scheuten were held in arenas without food at 95% RH and 20°C, the percentages of mites surviving to protonymphs were 5.0, 81.3, 86.3, and 83.8%, respectively. Unfed M. occidentalis larvae starved within 2–3 days, while immatures of the other three species lived up to 12–14 days, with some becoming adults by cannibalizing and/or scavenging. Phytosciid larvae given eggs, larvae/protochrysalis/protonymphs (L/P), deutochrysalis/deutonymphs (D) or teleiochrysalis/female adult (T/A) of Tetranychus urticae Koch, fed at different incidences during 6 h tests. Larvae of T. pyri never fed, but almost all larvae of M. occidentalis fed on eggs and L/Ps and 60–70% of M. occidentalis larvae fed on Ds and T/As. N. fallacis and A. andersoni larvae fed at incidences from 20–75% depending on the stage of spider mite given. Larvae fed more commonly on eggs and L/Ps than Ds and T/As for M. occidentalis and N. fallacis but not A. andersoni. Protonymphs and deutonymphs of all four species, readily fed on T/As after 3 h of exposure, but incidences were higher for A. andersoni and T. pyri. Feeding on phytoseiid larvae by protonymphs and deutonymphs also was more common for A. andersoni and T. pyri. Except for M. occidentalis, deutonymphs fed more than protonymphs on phytoseiid larvae. Results are discussed in relation to individual species life histories and the value of these traits in predicting a species role in a biological control system.     

Effects of humidity on eggs and immatures of Neoseiulus fallacis,Amblysieus andersoni,Metaseiulus occidentalis and Typhlodromus pyri (Phytoseiidae): implications for biological control on apple,caneberry, strawberry and hop

The lethal humidity (LH₅₀) responses at 20°C of eggs of two strains of Neoseiulus fallacis (Garman) were 71.6 and 69.7%; of three strains of Amblyseius andersoni (Chant) were 62.9, 62.0 and 62.4% and of one strain each of Typhlodromus pyri Scheuten and Metaseiulus occidentalis Nesbitt were 55.0 and 28.4%, respectively. Eggs of three genetically distinct strains of A. andersoni from Oregon, the Netherlands and Italy did not respond differently from one another nor did eggs of freely hybridizing N. fallacis from Michigan and Oregon. Mortality of larvae through development to early protonymphs at 50% RH, 20°C. was 91.9, 82.3, 46.2 and 31.0% for fed mites and 98.1, 83.2, 67.0 and 89.7% for unfed mites of Oregon strains of N. fallacis, A. andersoni, T. pyri and M. occidentalis, repectively. Fed larvae-protonymphs of M. occidentalis and T. pyri were more tolerant of low humidity than fed larvae-protonymphs of N. fallacis and A. andersoni. Mortality was less for fed than unfed larvae-protonymphs of M. occidentalis and T. pyri, but there were no differences for A. andersoni and N. fallacis. Levels of feeding by predator larvae on T. urticae and cannibalism by phytoseiid protonymphs contributed to species differences. Responses to humidity are discussed in relation to geographic and host plant distributions and biological control by single or mixed species populations of phytoseiids.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).     

Absorption and cannibalism: do phytoseiids conserve egg resources when prey densities decline rapidly?

The number of eggs oviposited or left in the opisthosomas of dead mites (total eggs) was assessed for Metaseiulus occidentalis (Nesbitt), Neoseiulus fallacis (Garman), Typhlodromus pyri Scheuten or Amblyseius andersoni Chant when each was caged with either (1) no Tetranychus urticae Koch, (2) only odours of T. urticae, (3) ten eggs of M. occidentalis or (4) ten nymphs of M. occidentalis (T. pyri for M. occidentalis). The total eggs for the no prey versus odour tests did not differ within species; the levels were the greatest for N. fallacis > T. pyri > A. andersoni > M. occidentalis. Among treatments, egg means did not differ for M. occidentalis but they did for N. fallacis and T. pyri and similar trends were seen for A. andersoni. Egg means were usually less for mites held with ten predator nymphs than mites held with ten predator eggs or with no prey. Were adult females with nymphs absorbing rather than ovipositing their eggs or dying with them in their opisthosomas? Activity levels (walking) for adult females were no more for mites held with nymphs versus no food. The data indicated that interference by nymphs was not increasing the energy use of females and thus reducing egg levels. However, tests with ten nymphs, one egg and no adult female had egg losses from nymphal predation that could account for fewer eggs in cage tests. Overall, no evidence for absorption was found. If it occurs, it must be among younger eggs or mites exposed to less rapid prey losses than were the mites tested here; in addition, other stimuli may cause absorption. The total eggs in sticky-tape tests were greatest for N. fallacis > M. occidentalis > T. pyri > A. andersoni. Cage versus stick-tape data differed most for M. occidentalis because of cannibalism. All four mites cannibalized eggs but M. occidentalis did most rapidly and extensively. When starved, it laid all of its eggs before the other three species did. Such behaviours may enhance survival of M. occidentalis when prey become scarce.     

Spatial scale of aggregation in three acarine predator species with different degrees of polyphagy

Aggregative responses by the predatory mites, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to spatial variation in the density of mobile stages of Tetranychus urticae (Acari: Tetranychidae) were studied over different spatial scales on greenhouse roses. Significant spatial variations in prey numbers per leaflet, per leaf, per branch or per plant were present in all experimental plots. None of the predator species responded to prey numbers per plant, and all searched randomly among plants. Within a plant, the oligophagous P. persimilis searched randomly among branches, but aggregated strongly among leaves within a branch and among leaflets within a leaf. The narrowly polyphagous T. occidentalis searched randomly among leaflets within a leaf and amond leaves within a branch, but aggregated strongly among leaflets or leaves within a plant. The boradly polyphagous A. andersoni searched randomly among leaflets within a leaf, a branch or a plant, and among leaves within a branch or a plant, but distributed themselves more often on branches with lower prey densities. Thus, specialist predators aggregate strongly at lower spatial levels but show random search at higher spatial levels, whereas generalist predators show random search at lower spatial levels but aggregate at higher spatial levels. This is the first empirical evidence demonstrating the relation between the degree of polyphagy and the spatial scale of aggregation. It is also concluded that both the prey patch size (i.e. grain) and predator foraging range (i.e. extent) are important for analyzing spatial scales of predator aggregation. The importance of studying spatial scale of aggregation is also discussed in relation to predator-prey metapopulation dynamics.     

Different colonization patterns of phytophagous and predatory mites (Acari: Tetranychidae, Phytoseiidae) on three grape varieties: a case study

In a vineyard having three varieties of grape (Merlot, Trebbiano and Garganega) differently colonized by two phytoseiid species,Typhlodromus pyri Scheuten andAmblyseius andersoni (Chant), the dynamics of mite populations were monitored over 5 years (1989–1993) in order to study their colonization, interspecific competition and the control of spider mites, i.e.Panonychus ulmi (Koch). These aspects were also investigated by releasingT. pyri, A. andersoni andAmblyseius aberrans (Oudemans) on some of the above varieties. In most of the experimental years (1989–1992), selective pesticides were used in order to allow a successful release of phytoseiids, in particularA. aberrans. The use of non-selective insecticides was re-established during 1993 in order to test its effect on the new mite communities originating from 1989 onwards. In the first years of the experiments an apparent relationship between grape variety and phytoseiid species was observed: in the control plots,A. andersoni occurred on Merlot whereT. pyri was rare, while the latter species was largely dominant overA. andersoni on Trebbiano and Garganega.Panonychus ulmi populations reached moderate levels only on Merlot and in the first part of experiments. The variety-phytoseiid species relationship was temporary as, at the end of experiments,T. pyri was completely dominant on all varieties. This new situation started when prey occurrence and interspecific competition decreased in importance. The moderate success of theT. pyri release on Merlot contrasts with the results of previous experiments. Two factors could be involved in this phenomenon: low interspecific competition by phytoseiids and predation by macropredators.Amblyseius aberrans was able to displaceA. andersoni andT. pyri on grape varieties where the two species were more abundant and reached higher population densities on varieties with pubescent leaf undersurfaces. In the first experimental year, spider mite densities were reduced more effectively inA. aberrans release plots than in the control or inT. pyri release plots. One year later,P. ulmi reached lower levels in the release treatments than in the control.Typhlodromus pyri andA. aberrans persisted in conditions of prey scarcity. The high competitivity ofA. aberrans over the remaining two phytoseiid species constitutes a major factor in selecting predatory species for inoculative releases in vineyards.     

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 Typhlodromus 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. Predator–prey 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.     

Interactions among predators and phytophagous mites on apple; possible impact on biocontrol of Panonychus ulmi by Typhlodromus pyri in orchards

Laboratory experiments were conducted to determine the potential impact of the phytoseiid Euseius finlandicus, the mirid Blepharidopterus angulatus and the anthocorid Orius majusculus on the Typhlodromus pyri/Panonychus ulmi predator/prey relationship on apple. Euseius finlandicus consumed more immature spider mites than did T. pyri. When both phytoseiids were present and spider mite prey was abundant, there was no evidence of a negative interaction between the predators. In experiments where each predatory mite was confined with large numbers of the other predator, interspecific predation was exhibited by adults of each species on immatures of the other, but more so by E. finlandicus. In the predatory insect/phytoseiid experiments, when confined with spider mites and large numbers of T. pyri, both B. angulatus and O. majusculus consumed some T. pyri, but spider mites were the preferred prey. In experiments with B. angulatus, O. majusculus and T. pyri feeding on P. ulmi, there was no evidence of negative interactions between the predatory insects and T. pyri.     

The Transfer of Typhlodromus pyri on Grape Leaves for Biological Control of Panonychus ulmi (Acari: Phytoseiidae, Tetranychidae) in Vineyards in Ontario, Canada

The phytophagous mite Panonychus ulmi Koch has become a significant problem in Ontario vineyards. We attempted to introduce and establish populations of the predatory mite Typhlodromus pyri Scheuten for P. ulmi biological control. Grape leaves were transferred from a vineyard containing T. pyri in early summer 1998, by picking leaves from a donor vineyard and attaching them onto leaves in the release vineyard where T. pyri were extremely rare. Two release treatments were used with rates of 8.5 (1×) and 25.5 (3×) mobiles per vine. In the first season, T. pyri established in similar densities in both release treatments, which were significantly higher than controls. However, there were no differences among treatments in P. ulmi densities in 1998 as a result of predator release. During summer 1999, significantly fewer P. ulmi mite-days were observed in release plots compared to the control. Amblyseius fallacis (Garman) was common throughout the release vineyard in 1998 and in 1999, but appeared on the vines too late in the season to maintain low P. ulmi densities. T. pyri appeared to out-compete A. fallacis in 1999 because A. fallacis densities were significantly lower in plots where T. pyri had been released than in control plots. We conclude that T. pyri can be effective for P. ulmi biological control in Ontario vineyards and may be introduced by transferring leaves. In Europe, transferring prunings has been the standard method of inoculating T. pyri into new vineyards. Here we show that transferring leaves is another practical method.     

Developmental times and oviposition rates of predatory mitesTyphlodromus pyri andAmblyseius andersoni (Acari: Phytoseiidae) reared on different foods

The developmental times and the reproduction of two resistant Italian strains ofTyphlodromus pyri Scheuten andAmblyseius andersoni (Chant) were studied in the laboratory by rearing them on the spider mitesPanonychus ulmi (Koch) andEotetranychus carpini (Oud.), on the eriophyidColomerus vitis (Pgst.) and on pollen ofMesembryanthemum criniflorum. The response ofT. pyri andA. andersoni females to a spider mite supply (P. ulmi orE. carpini) of 4, 8 and 16 adult female prey per female predator per day was also studied.Development ofT. pyri onE. carpini andC. vitis required a shorter period than onM. criniflorum pollen, while intermediate values were recorded forP. ulmi. When the highest number of prey was offered, the influence of different foods on oviposition rates ofT. pyri was not significant. An increase in spider mite supply favoured a shorter pre-oviposition period and higher oviposition rates.Development ofA. andersoni was faster on pollen than on spider mites, while intermediate values were found concerningC. vitis. Differences statistically significant were recorded for development onP. ulmi andC. vitis. Colomerus vitis proved to be the more suitable food in terms of oviposition. The oviposition rate decreased when feeding uponP. ulmi, but reached intermediate values onE. carpini andM. criniflorum. Increasing spider mite densities caused shorter pre-oviposition times and higher oviposition rates. Using a given number ofE. carpini females, rather than those ofP. ulmi, resulted in higher oviposition rates and shorter pre-oviposition times.For both predators, the results suggest a higher intrinsic rate of population increase onE. carpini orC. vitis than onP. ulmi.The research was supported by a grant from Regione Veneto (Lotta biologica ed integrata nel controllo di insetti ed acari dannosi).The general lines of the research have been planned by C. Duso. The authors contributed equally to the experimental work.     

Comparison of the responses of two predaceous mites,Typhlodromus pyri and Zetzellia mali,to variation in prey density

The ability of a predator to respond to prey density in a patchy habitat has been the focus of much study in biological control systems and elsewhere. Here we look at the response of two species of predatory mite commonly found in commercial apple orchards, Typhlodromus pyri Scheuten and Zetzellia mali Ewing. The two species differ in several characteristics: T. pyri is a more successful biological control agent, is more mobile, has a slightly narrower breadth of diet and prefers the target prey, Panonychus ulmi Koch. We measured resisdence time as a function of prey density, both under field and laboratory conditions. Both predators showed a wide variation in behaviour; however, Z. mali increased residence time in response to the presence of prey, while T. pyri did not show a significant response. Both predators usually left a patch before all prey were consumed. Patterns from the field and laboratory were similar, although residence time was shorter under field conditions.     

Toxicity of a number of pesticides to strains ofTyphlodromus pyri andAmblyseius andersoni (Acari: Phytoseiidae)

Side effects of ten pesticides used in orchards and vineyards were tested with a laboratory method on several Dutch and Italian strains of the predatory mitesTyphlodromus pyri Scheuten andAmblyseius andersoni (Chant). Resistant and susceptible strains of both species were studied. Results showed that a test which evaluates mortality of various developmental stages and fecundity of adult females is better than one that measures only survival of adult females. A definite resistance to certain pesticides was found in ItalianT. pyri andA. andersoni. The level of resistance to parathion, azinphos-methyl and carbaryl was particularly high in some strains ofA. andersoni. The high level of resistance to certain pesticides was often associated with a marked reduction in fecundity.      

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.     

Establishing insecticide-resistant phytoseiid mite predators in deciduous tree fruit orchards

A summary of basic biological, ecological and toxicological characteristics of 3 phytoseiid mite predatorsTyphlodromus occidentalis Nesbitt,Amblyseius fallacis Garman andTyphlodromus pyri Scheuten which have developed resistances to certain insecticides commonly used in commercial fruit orchards is presented. Methods of culturing and mass-rearing these mites are reviewed. Also the cultural and pesticidal measures which should be maintained to facilitate effective biological control of spider mite pest of deciduous fruit crops by these predators and to maximally provide for their establishment in areas of the world where resistance has not developed or they do not occur naturally are discussed.

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Résumé L'auteur résume les caractéristiques fondamentles biologiques, écologiques et toxicologiques de 3 phytoséiides prédateurs d'acariens,Typhlodromus occidentalis Nesbitt,Amblyseius fallacis Garman etTyphlodromus pyri Scheuter, qui ont acquis une résistance à certains insecticides généralement utilisés dans les vergers de fruits commerciaux. Les méthodes d'élevage et de multiplication massive de ces acariens sont rappelées. Sont également discutées les techniques culturales et phytosanitaires qui sont à employer pour favoriser ces prédateurs comme agents de lutte biologique efficace contre les acariens nuisibles aux vergers et pour procéder à leur implantation dans les régions du monde où la résistance ne s'est pas manifestée ou bien où ces prédateurs n'existent pas naturellement.

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This publication was supported in part by a NSF-EPA grant GB-34718 to the University of California. The findings, opinions and recommendations expressed herein are those of the author and not necessarily those of the University of California, NSF or EPA. Published as Journal Article N. 1559 of the Michigan State University Agricultural Experiment Station.     

Analysis of prey preference in phytoseiid mites by using an olfactometer,predation models and electrophoresis

Prey preference of three phytoseiid species,Typhlodromus pyri Scheuten,Amblyseius potentillae (Garman) andA. finlandicus (Oudemans) which occur in Dutch orchards, was analysed with respect to two economically important phytophagous mites, the European red spider mitePanonychus ulmi (Koch), and the apple rust miteAculus schlechtendali (Nalepa). Two types of laboratory experiments were carried out: (1) olfactometer tests to study the response when volatile kairomones of both prey species were offered simultaneously; and (2) predation tests in mixtures of the two prey species and comparison with calculated predation rates, using a model provided with parameters estimated from experiments with each prey species alone. In addition, the diet of field-collected predators was analysed using electrophoresis. For each predator species the results of the different tests were consistent, in thatT. pyri andA. potentillae preferredP. ulmi overA. schlechtendali, whereasA. finlandicus preferredA. schlechtendali overP. ulmi.