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.     

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.     

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.     

Adaptability and efficacy of transgenic and wild-type Metaseiulus occidentalis (Acari: Phytoseiidae) compared as part of a risk assessment

Comparisons between transgenic (T) and wild-type Metaseiulus occidentalis colonies (COS) were made under laboratory conditions as part of a risk assessment effort prior to proposed field releases. There were no differences between the transgenic T18 colony and the COS strain in the daily egg production, hatchability at three temperatures and four relative humidity (RH) conditions, diapause incidence, or proportion of female progeny produced. Metaseiulus occidentalis eggs do not hatch at 38°C under any RH tested, nor at 33.5°C under 100% RH, indicating that high temperature and extreme RH affect egg hatch negatively. At 28.5 and 33.5°C, fewer eggs hatched at 32.5% than at 75.5 and 93% RH. Metaseiulus occidentalis cannot survive on any plant tested without prey nor on a diet of pollen alone; adult female mites cannot prey or survive on a diet of eggs and larvae of two lepidopteran species, indicating that the suitability of food sources has not been altered in the transgenic strain. Two subcolonies, derived from two transgenic strains using single females, differed in the rate of egg hatch at 28.5°C under 32.5 and 100% RH, indicating that reduced genetic variation and/or random genetic drift in the two lines may have led to differences in some biological characters. Since we did not find any significant differences between the T18 and COS colonies in the traits tested, the T18 colony is not expected to exhibit any new biological attributes in a proposed short-term field release.     

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.     

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.     

Factors affecting biological control of apple mites by mixed populations ofMetaseiulus occidentalis andTyphlodromus pyri

Metaseiulus occidentalis (Nesbitt) andTyphlodromus pyri Scheuten have complementary features/ traits that enable them to control effectively plant-feeding mites on apple. Populations of both predators gave as good or better biological control of the apple rust mite (Aculus schlechtendali Nalepa), European red mite (Panonychus ulmi Koch) and two-spotted spider mite (Tetranychus urticae (Koch)) than single-predator populations. With mixed predators,M. occidentalis provided better control of spider mites the first season after release, butT. pyri gave better control in the second season. Several factors affected the ability of predators to provide biological control: When prey were dense,M. occidentalis rapidly increased during the warm mid-summer, whileT. pyri provided greater predation when it was cool at the start or end of the growing season. When few prey were present, searching byM. occidentalis was more confined on individual apple leaves, but it migrated between leaves and trees more often. Pollen feeding, cannibalism and interspecific predation were more common byT. pyri. In fall, oviposition byM. occidentalis stopped sooner and in the following spring,T. pyri reproduced beforeM. occidentalis. Research needs and management of mixed-predator populations are discussed.     

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.     

Effect of prey density on sex ratio of two predacious mites, Phytoseiulus persimilis and Amblyseius womersleyi (Acari: Phytoseiidae)

The sex ratios of two phytoseiid mites, Phytoseiulus persimilis and Amblyseius womersleyi, were observed under various prey conditions. Upon consumption of abundant prey, both phytoseiids produced progeny in a female-biased sex ratio (approximately 0.8 females). When few prey were consumed, the sex ratio was lowered to 0.5 (the unbiased sex ratio). Under the conditions in which the unbiased sex ratio was observed, male and female progeny appeared in an alternating sequence. To determine the change in the sex ratio and the sequence of progeny, the size of eggs deposited by females under various prey conditions was first compared. Survivorship and developmental rate of progeny (immatures) hatched under ample and poor prey conditions were also examined. The eggs deposited under poor prey conditions were smaller than those deposited under ample prey conditions. This is an indication that the phytoseiid females did not invest extra energy into the eggs to secure survival of their progeny under poor prey conditions. The male and female progeny from the small eggs developed slowly, probably due to the small egg size. However, hatchability of the small eggs and survival of the immatures were the same as those of the normal eggs when the immatures were reared under ample prey conditions. The immature survivorship was little affected by the prey consumption rate of their mothers when the immatures were reared under poor prey conditions. We concluded that the sex ratio of phytoseiid mites is not determined by the characteristics of the progeny, but by the nutritious condition of the females.Exp Appl Acarol 22: 709723 © 1998 Kluwer Academic Publishers     

Transmission of injected DNA sequences to multiple eggs of Metaseiulus occidentalis and Amblyseius finlandicus (Acari: Phytoseiidae) following maternal microinjection

The persistence of DNA injected into two species of adult female phytoseiids and its transmission to serial eggs deposited by them was assessed by the polymerase chain reaction (PCR). The effect of DNA concentration on persistence and transmission was examined in Metaseiulus occidentalis. M. occidentalis females were microinjected with plasmid DNA at three different concentrations (250, 500, 750 ng L^–1) and allowed to deposit one to five eggs before the females and their last eggs were analyzed. Plasmid DNA was found in 82% of the females assayed and in 70% of all the eggs analyzed (including the fifth eggs produced after microinjection). Transmission of DNA to multiple eggs was also examined in Amblyseius finlandicus. Females of this species are less traumatized by microinjection allowing analysis of transmission over a more extended number of eggs. Females were microinjected and allowed to deposit eggs until their death. DNA from every fifth egg was analyzed by the PCR. PCR products were amplified from 51% of the eggs and from all egg classes except the 30th egg. The persistence and presence of plasmid DNA in both eggs and females suggests that (1) maternal microinjection is a more efficient method for DNA delivery than traditional egg microinjection, (2) it may be possible to isolate transformants from fewer maternally-microinjected females than originally expected, and (3) maternal microinjection could be useful as a DNA delivery system in other phytoseiids.     

Evaluation of the predacious mite Hemicheyletia wellsina (Acari: Cheyletidae) as a predator of arthropod pests of orchids

The cheyletid predator Hemicheyletia wellsina was found in association with orchid pests in a Florida greenhouse. The life history of H. wellsina was determined using the two-spotted spider mite, Tetranychus urticae, as prey in the laboratory. Hemicheyletia wellsina is arrhenotokous, with mated females producing a female-biased sex ratio of 3.4:1. Adult longevity averaged 30.2 days, with females producing an average of 18.5 eggs. Adult females live up to 17 days without food, surviving an average of 9.7 days. Hemicheyletia wellsina will readily feed on the phytoseiid Metaseiulus occidentalis, suggesting intraguild predation by H. wellsina on phytoseiids could be important in pest management programs. The average prey consumption of H. wellsina is 1.5 T. urticae females per day. The results of this study provide information on an under-studied group of mite predators. Although H. wellsina does not have the qualities that would be necessary for its use in an augmentative or classical biological control program, its use in naturally occurring biological control could be beneficial.     

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; tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, N. 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.     

Interspecific competition and predation between immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Interspecific competition and predation in immature Amblyseius fallacis (Garman), Amblyseius andersoni Chant, Typhlodromus occidentalis (Nesbitt) and Typhlodromus pyri Scheuten were examined in small cages at three egg densities (0, 20 and 80) of two-spotted spider mite, Tetranychus urticae Koch, in the laboratory at 25±1°C,80% RH and 16L: 8D photoperiod. For the six possible between-species comparisons, the large polyphagous A. andersoni always outcompeted the other three predator species, which were either smaller and/or less polyphagous; the small oligophagous T. occidentalis was always eliminated by the other three predator species, which were either larger and/or more polyphagous. The small and polyphagous T. pyri tied with the large and oligophagous A. fallacis. The outcome of the interaction was generally similar at the three prey densities except in (1) the A. fallacis-A. andersoni system where the advantage of A. andersoni over A. fallacis was reduced when 20 or 80 eggs per cage were present at the start of the interaction and (2) the A. fallacis-T. occidentalis system where the advantage of A. fallacis over T. occidentalis increased with prey density. This study indicates that predator size, predator degree of polyphagy and prey density can affect the competitiveness of immature phytoseiids.     

Dispersal between and colonization of apple byMetaseiulus occidentalis andTyphlodromus pyri (Acarina: Phytoseiidae)

Adult femaleMetaseiulus occidentalis Nesbitt immigrated faster and at higher rates than didTyphlodromus pyri Scheuten to mini-orchards of young apple trees at 0, 10 and 100 m downwind from four apple orchards in the Hood River Valley, Oregon. Colonization of mini-orchards also was faster byM. occidentalis. Rates of emigration were similar between species as measured by movement into trees placed within, but not in contact with source orchard trees. Dispersal and colonization may be influenced by intrinsic physiological and morphological attributes, food habits, and responses to prey and habitat requirements. Implications of dispersal and colonization by these mites to biological control and pesticide resistance management are discussed.     

Influence of food and temperature on development and oviposition ofEuseius stipulatus andTyphlodromus phialatus (Acari: Phytoseiidae)

The influence of feeding and temperature on life table parameters ofEuseius stipulatus (Athias-Henriot) andTyphlodromus phialatus Athias-Henriot was determined in laboratory tests. Both phytoseiids developed and reproduced readily when fed onCarpobrotus edulis (L.) pollen andPanonychus citri (McGregor) Intrinsic rates of increase (r _m ) at 25°C forE. stipulatus andT. phialatus were 0.197 and 0.126, and, when fed onP. citri, 0.129 and 0.144, respectively.Tetranychus urticae Koch offered as prey allowed normal development and oviposition ofT. phialatus, but no eggs were laid byE. stipulatus when fed on this spider mite. Eggs and honeydew ofAleurothrixus floccosus Mask. furnished sufficient nutrition for development of immatures ofE. stipulatus. Larvae ofPlanoccocus citri Risso, and honeydew and eggs ofA. floccosus, allowed adult survival but no egg-laying of the predaceous mites.Lorryia formosa Cooreman was not a favoured prey species.Life tables were calculated forE. stipulatus fed on pollen andT. phialatus fed onP. citri at constant temperatures of 18, 25, and 32°C. Maximum development was reached at 32°C, withr _m values of 0.225 forE. stipulatus and 0.179 forT. phialatus. In general, both phytoseiids showed medium to high total number of eggs per female and long oviposition periods when compared with other phytoseiid species.When mating took place at 32°C,E. stipulatus females were not able to lay eggs, thus suggesting an interference of high temperatures with fertilization in this species. No hatching was observed in eggs of either species when kept at relative humidities of 50% or lower. The possible significance of these responses is discussed in relation to population trends observed in the field.     

Intra- and interspecific predation by adult female Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) when provisioned with varying densities and ratios of Tetranychus urticae (Acari: Tetranychidae) and phytoseiid larvae

The phytoseiid mites Metaseiulus occidentalis (Nesbitt) and Typhlodromus pyri Schueten are used together and alone as biological control agents against tetranychid pest mites of apple. Their effectiveness as control agents may be impacted by intraguild predation. The effects of prey species and prey density on the rates of inter- and intraspecific predation and oviposition by these two predators were investigated through a series of experiments. Adult female predators were given prey as mixed populations of phytoseiid larvae and larvae of a more preferred species, the spider mite, Tetranychus urticae Koch, at different densities and ratios. Typhlodromus pyri, more of a generalist predator, showed higher rates of predation and cannibalism on phytoseiid immatures at most prey densities and ratios. Manly preference indices indicated that T. pyri switched to feed on phytoseiid larvae at higher prey levels and ratios of T. urticae than M. occidentalis. This greater ability to use phytoseiid larvae as prey may help stabilize T. pyri populations when more preferred prey is unavailable. This may, in part, explain the observed persistence of T. pyri populations when M. occidentalis populations were decreasing in orchard test plots.     

Distribution of the Predatory Mite Typhlodromus pyri (Acari: Phytoseiidae) on Different Apple Cultivars

The effect of apple cultivar on the distribution of the predatory mite Typhlodromus pyri was studied in an experimental orchard where spider mites occurred at negligible densities. Seven apple scab-resistant cultivars, showing some differences in their leaf morphology, were considered. In particular, their leaf blade was classified according to four levels of pubescence. The distribution of T. pyri along the shoots was also studied. In the first experimental year the colonization of different cultivars by T. pyri showed definite patterns, sometimes influenced by the occurrence of eriophyids. One year later, T. pyri abundance again showed some differences among cultivars in conditions of prey scarcity. In both years large phytoseiid populations were recorded on the cultivar N.Y. 18491, despite the low eriophyid occurrence, probably because of its highly pubescent leaf undersurfaces. In contrast, Prima and TSR 29T219, characterized by slightly pubescent leaf undersurfaces, supported low phytoseiid densities independently of prey availability.     

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.     

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.     

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).