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.     

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.     

Effects of prey mite species on life history of the phytoseiid predators Typhlodromalus manihoti and Typhlodromalus aripo

The effects of prey mite suitability on several demographic characteristics of phytoseiid predators and the relationship of these effects to the potential of phytoseiid predators to control herbivorous mite populations are well documented. Evidence has also accumulated in the last 20 years demonstrating that phytoseiid predators utilize herbivorous prey mite-induced plant volatiles as olfactory cues in locating their herbivorous mite prey. but less well established is the predictability of reproductive success from the ability of the predators to utilize olfactory cues to locate their prey, and how these processes are related to the success of the predators as biological control agents of the herbivorous mite. In this study, we determined in laboratory no choice experiments, the development, survivorship and fecundity of the two neotropical phytoseiid predators Typhlodromalus manihoti Moraes and T. aripo DeLeon when feeding on three herbivorous mites, including the key prey species Mononychellus tanajoa (Bondar), and the two alternative prey species Oligonychus gossypii (Zacher) and Tetranychus urticae (Koch). Intrinsic rate of increase (rm) of T. aripo was 2.1 fold higher on M. tanajoa as prey compared with T. urticae as prey, while it was almost nil on O. gossypii. For T. manihoti, rm was 2.3 fold higher on M. tanajoa as prey compared with O. gossypii as prey, while reproduction was nil on T. urticae. An independent experiment on odor-related prey preference of the two predator species (Gnanvossou et al. 2002) showed that T. manihoti and T. aripo preferred odors from M. tanajoa-infested leaves to odors from O. gossypii-infested leaves. Moreover, both predator species preferred odors from M. tanajoa-infested leaves over those from T. urticae-infested leaves. As reported here, life history of the two predatory mites matches odor-related prey preference if the key prey species is compared to the two inferior prey species. The implications of our findings for the persistence of T. manihoti and T. aripo and biological control of M. tanajoa in the cassava agroecosystem in Africa are discussed.     

Effects of fungicide residues on the survival, fecundity, and predation of the mites Tetranychus urticae (Acari: Tetranychidae) and Galendromus occidentalis (Acari: Phytoseiidae)

Representative fungicides from three or four families used for management of powdery mildew and other diseases in tree fruits were evaluated for their effects on a common spider mite and predator mite species, respectively. A modified Munger cell technique was effective in measuring the response of phytophagous and predaceous mites to fungicide residues on detached leaves in the laboratory. Demethylation-inhibiting (DMI) (imidazole [triflumazole] and triazole [myclobutanil]) and strobilurin (trifloxystrobin) fungicides were not toxic to female Tetranychus urticae Koch and Galendromus occidentalis (Nesbitt), and no sublethal effects were found on fecundity and predation rate after 3-5-d exposure to residues. Benomyl, a benzimidazole fungicide, increased adult mortality and reduced fecundity for both mite species; however, it did not alter the predation rate of G. occidentalis females on T. urticae eggs and larvae. Female G. occidentalis that survived the lethal effects of benomyl and the comparison acaricide pyridaben were unimpaired in predation. Our results for benomyl substantiate those of earlier studies and provide evidence for nontoxic effects of DMI and strobilurin fungicides on mites. We propose that DMI and strobilurin fungicides are a good fit for integrated mite management programs due to conservation of phytoseiid predatory mites.     

东亚小花蝽对西方花蓟马和二斑叶螨的捕食选择性

多食性天敌对猎物的捕食选择性及猎物密度对天敌捕食的干扰作用直接影响到天敌对不同猎物的控制作用.在实验室条件下研究了东亚小花蝽对西方花蓟马和/或二斑叶螨不同虫态的捕食选择性,以及两猎物中一种猎物的密度变化对小花蝽取食另一种猎物的影响.结果如下:东亚小花蝽5龄若虫和成虫对西方花蓟马2龄若虫的捕食选择性均强于对其成虫,对二斑叶螨的选择性为雌成螨>若螨>幼螨.实验中西方花蓟马2龄若虫是东亚小花蝽最喜好的虫态.二斑叶螨雌成螨密度固定为60头/19.63cm2,西方花蓟马若虫密度从10增加到60时显著地减少了东亚小花蝽对二斑叶螨的取食.反之,固定西方花蓟马同样密度,增加二斑叶螨密度却没有显著改变小花蝽对西方花蓟马的取食.此结果进一步表明,西方花蓟马是东亚小花蝽更喜好的猎物.     

Physiological performance of juvenile Haliotis rufescens and Haliotis discus hannai abalone exposed to the withering syndrome agent

The presence of Wolbachia and Cardinium bacteria has been documented in many arthropod species, including the predatory mite Metaseiulus (=Typhlodromus or Galendomus) occidentalis (Nesbitt) (Acari: Phytoseiidae). We show that Tetranychus urticae, the prey of Metaseiulus occidentalis, contains Wolbachia and no detectable Cardinium using quantitative PCR (qPCR). Starvation for 72 h at 22°C eliminated most, if not all, Wolbachia in M. occidentalis adult females from 7 laboratory colonies. Refeeding of M. occidentalis with T. urticae after starvation for 72 h restored the amounts of Wolbachia in M. occidentalis to those of prestarvation levels, suggesting that Wolbachia detected in M. occidentalis starved for shorter periods of time in current, and some previous, studies likely came from T. urticae. Furthermore, eggs from all M. occidentalis colonies examined were free of Wolbachia if they were surface-decontaminated with 0.3% sodium hypochlorite before DNA extraction. Cardinium was present in 6 of 14 laboratory colonies of M. occidentalis. Starvation for 3, 24, 48, and 72 h had no effect on the amounts of Cardinium in adult females from the Cardinium-positive colonies. Eggs from these colonies were positive for Cardinium but contained less than 1% of the titers found in adult females. The data suggest that Cardinium, but not Wolbachia, is an endosymbiont in certain populations of M. occidentalis. In light of our current findings, we recommend specific practices for the identification of potential symbionts in predatory arthropod species using the PCR.     

Verified and potential pathogens of predatory mites (Acari: Phytoseiidae)

Several species of phytoseiid mites (Acari: Phytoseiidae), including species of the genera Amblyseius, Galendromus, Metaseiulus, Neoseiulus, Phytoseiulus and Typhlodromus, are currently reared for biological control of various crop pests and/or as model organisms for the study of predator-prey interactions. Pathogen-free phytoseiid mites are important to obtain high efficacy in biological pest control and to get reliable data in mite research, as pathogens may affect the performance of their host or alter their reproduction and behaviour. Potential and verified pathogens have been reported for phytoseiid mites during the past 25 years. The present review provides an overview, including potential pathogens with unknown host effects (17 reports), endosymbiotic Wolbachia (seven reports), other bacteria (including Cardinium and Spiroplasma) (four reports), cases of unidentified diseases (three reports) and cases of verified pathogens (six reports). From the latter group four reports refer to Microsporidia, one to a fungus and one to a bacterium. Only five entities have been studied in detail, including Wolbachia infecting seven predatory mite species, other endosymbiotic bacteria infecting Metaseiulus (Galendromus, Typhlodromus) occidentalis (Nesbitt), the bacterium Acaricomes phytoseiuli infecting Phytoseiulus persimilis Athias-Henriot, the microsporidium Microsporidium phytoseiuli infecting P. persimilis and the microsporidium Oligosproridium occidentalis infecting M. occidentalis. In four cases (Wolbachia, A. phytoseiuli, M. phytoseiuli and O. occidentalis) an infection may be connected with fitness costs of the host. Moreover, infection is not always readily visible as no obvious gross symptoms are present. Monitoring of these entities on a routine and continuous basis should therefore get more attention, especially in commercial mass-production. Special attention should be paid to field-collected mites before introduction into the laboratory or mass rearing, and to mites that are exchanged among rearing facilities. However, at present general pathogen monitoring is not yet practical as effects of many entities are unknown. More research effort is needed concerning verified and potential pathogens of commercially reared arthropods and those used as model organisms in research.     

Consumption rate, functional response and preference of the predaceous mite Iphiseius degenerans to Tetranychus urticae and Eutetranychus orientalis

The functional response of females of the phytoseiid mite, Iphiseius degenerans (Berlese), to increasing densities of females of its prey, Tetranychus urticae Koch and Eutetranychus orientalis Klein, on bean leaves, were studied under laboratory conditions. Our results indicated that the predator consumed significantly more items of E. orientalis than of T. urticae at all densities treatments. Daily consumption of the predator increased with increasing prey density until a plateau was reached-maximum number of prey consumed was ca. 4 for T. urticae and ca. 12 for E. orientalis. A Type II functional response was determined by a logistic regression model. The highest estimated value a (instantaneous rate of attack) and the lowest value of T ( h ) (handling time) were found for the predator feeding on E. orientalis. Prey selection was evaluated by simultaneously presenting both prey species to the predator in various ratios and at increasing densities. I. degenerans showed a higher predation rate and higher preference for E. orientalis at all the ratios and prey densities tested. This may be due to the smaller size or the inactivity of E. orientalis and the inability of the predator to cope with the webbing of T. urticae. Our results suggest that I. degenerans can be considered a suitable biological control candidate based on its preference for E. orientalis in the Mediterranean region.     

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.     

Effects of prey mite species on life history of the phytoseiid predators Typhlodromalus manihoti and Typhlodromalus aripo

The effects of prey mite suitability on several demographic characteristics of phytoseiid predators and the relationship of these effects to the potential of phytoseiid predators to control herbivorous mite populations are well documented. Evidence has also accumulated in the last 20 years demonstrating that phytoseiid predators utilize herbivorous prey mite-induced plant volatiles as olfactory cues in locating their herbivorous mite prey, but less well established is the predictability of reproductive success from the ability of the predators to utilize olfactory cues to locate their prey, and how these processes are related to the success of the predators as biological control agents of the herbivorous mite. In this study, we determined in laboratory no choice experiments, the development, survivorship and fecundity of the two neotropical phytoseiid predators Typhlodromalus manihoti Moraes and T. aripo DeLeon when feeding on three herbivorous mites, including the key prey species Mononychellus tanajoa (Bondar), and the two alternative prey species Oligonychus gossypii (Zacher) and Tetranychus urticae (Koch). Intrinsic rate of increase (r_m) of T. aripo was 2.1 fold higher on M. tanajoa as prey compared with T. urticae as prey, while it was almost nil on O. gossypii. For T. manihoti, r_m was 2.3 fold higher on M. tanajoa as prey compared with O. gossypii as prey, while reproduction was nil on T. urticae. An independent experiment on odor-related prey preference of the two predator species showed that T. manihoti and T. aripo preferred odors from M. tanajoa-infested leaves to odors from O. gossypii-infested leaves. Moreover, both predator species preferred odors from M. tanajoa-infested leaves over those from T. urticae-infested leaves. As reported here, life history of the two predatory mites matches odor-related prey preference if the key prey species is compared to the two inferior prey species. The implications of our findings for the persistence of T. manihoti and T. aripo and biological control of M. tanajoa in the cassava agroecosystem in Africa are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population growth and persistence when prey is diminishing in single-species and two-species systems of the predatory mites Euseius finlandicus, Typhlodromus pyri and Kampimodromus aberrans

Population growth and persistence of Euseius finlandicus (Oudemans), Typhlodromus pyri Scheuten and Kampimodromus aberrans (Oudemans) (Acari: Phytoseiidae) were studied in single-species and two-species systems on apple seedlings primarily infested by Panonychus ulmi Koch (Acari: Tetranychidae) in an environmentally controlled greenhouse. During the experiment, the seedlings developed natural infestations by Tetranychus urticae Koch (Acari: Tetranychidae), Frankliniella occidentalis Pergande (Thysanoptera: Thripidae), and powdery mildew. Several weeks after the start of the experiment a condition of diminishing prey availability was created by use of hexythiazox treatments. Without heterospecific competitors, T. pyri attained a higher population level than E. finlandicus or K. aberrans when similar amounts of food (spider mites) were available to each. Population growth of T. pyri was decisively favoured by the presence of T. urticae. In the single-species systems each predatory species persisted to the end of the experiment in spite of diminishing prey. In two-species systems with T. pyri/E. finlandicus and T. pyri/K. aberrans that were started with the same number of individuals of each species, only T. pyri was left at the end of the experiment. Typhlodromus pyri became more numerous than the other species when prey was abundant (which was in accordance with the results of the single-species groups) and finally displaced E. finlandicus and K. aberrans towards the end of the experiment. The following factors may have contributed to the dominance of T. pyri: (1) the ability of adult females to survive longer without food than those of E. finlandicus and K. aberrans, (2) the ability to complete juvenile development and to sustain reproduction with phytoseiid prey and (3) an advantage in foraging behaviour over K. aberrans and E. finlandicus at low spider mite levels. Euseius finlandicus predominated in the two-species system E. finlandicus/K. aberrans, but both species persisted to the end of the experiment.     

Prey preference of Orius sauteri between Western Flower Thrips and spider mites

Prey preference of polyphagous predators plays an important role in the suppression of various species of pest insects. In this study, the prey preference of Orius sauteri (Poppius) (Heteroptera: Anthocoridae) between deutonymphs of Tetranychus urticae Koch (Acari: Tetranychidae) and second instars of Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and the influence of spider mite density on the preference were examined in the laboratory in three-dimensional set-ups at 25 ± 1 °C and 70 ± 5% r.h. The results showed that the predatory bug in the presence of equal densities of the two prey species had a clear preference for thrips (preference index β: 0.86 ± 0.02), consuming 7.2 thrips larvae and 1.5 spider mite deutonymphs during the experimental periods of 6 h. The number of thrips consumed by O. sauteri did not change when the density ratio of thrips to spider mites was decreased from 1:1 to 1:5. Predation on spider mites increased when their ratio to thrips increased, but the disproportionate predation (i.e., the functional response) of O. sauteri towards spider mites resulted in a linear increase in the preference for F. occidentalis . The possible implications of these findings for the suppression of spider mites by O. sauteri in relation to the recent invasion of F. occidentalis into China are discussed.     

Spatial distribution and coincidence of Neoseiulus californicus and Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae) on strawberry

In strawberry greenhouses in La Plata (Buenos Aires, Argentina), Tetranychus urticae is a major pest and Neoseiulus californicus is its most important established phytoseiid predator. The purpose of this study was to determine the spatial distribution and coincidence of N. californicus and T. urticae on strawberry. T. urticae populations exhibited density variations that were followed, with some temporal delay, by those of the predator. In general terms, N. californicus exhibited lower aggregation than T. urticae. The index of dispersion (I) of T. urticae had values significantly greater than 1 in 100% of the sampling dates. The percentage of infested leaflets increased with prey density in a curvilinear way, ranging from 80 to 100% when density was higher than 100 individuals/leaflet. N. californicus had values of I higher than unity in 86.7% (G1) and 53.8% (G2) of the cases. TaylorÕs b and IwaoÕs values were greater than 1 for both populations, this indicating aggregation. IwaoÕs results showed larger aggregation units of T. urticae (=53.24) than of N. californicus (=3.61), probably due to its higher fecundity and oviposition behaviour of laying eggs in clumps. Both populations were overdispersed, although in general terms, N. californicus exhibited lower aggregation than T. urticae. This would create refuges for the prey, thus increasing the persistence of the system. Index of coincidence of predator (Ic) was high most of the time even at low densities of both populations, decreased abruptly at the end of an interaction cycle and was similar at greenhouse and plant spatial scales. The high spatial coincidence of N. californicus with T. urticae suggests an important dispersal capacity of the predator and a high ability to detect leaflets with prey. The refuge index for the prey decreased with predator density and was higher at greenhouse scale than at plant scale, both at similar predator densities. At greenhouse scale and despite high predator density, the prey had a 10%-refuge, which would lead to the system persistence. Results of the present study suggest that N. californicus is a promising established natural enemy for controlling T. urticae on strawberry.     

Predation on two vole species by a shared predator: antipredatory response and prey preference

In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitat-specific characteristics of these prey species are playing a minor role compared to sex-specific characteristics.     

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.     

Classifying Life-Style Types of Phytoseiid Mites:Diagnostic Traits

Several traits are useful for identifying life-style types of predaceous phytoseiid mites when either 2 (diet generalist-specialist) or 4 (specialist I and II-generalist III and IV) type models [McMurtry J.A. and Croft B.A. 1997. Annu. Rev. Entomol. 42: 291-321] are considered. Traits useful for both models are developmental time and oviposition rates when feeding on several food types. Discriminating for the 2-types model are dorsal shield setae lengths, and intra- and inter-specific predation. Another trait useful for both models is feeding preferences of adult female phytoseiids on eggs versus larvae of Tetranychus urticae Koch. In this paper, we review established and other traits that need more study such as mouthpart types, other morphological features, spider mite webbing effects, distributions relative to prey-foods, plant-host relationships including domatia and sap feeding, density-dependent responses to prey and predator-prey ratios required for biological control. Uses of life-style data in biological control decision-making are discussed.     

Prey Selection by the Lady Beetle Harmonia axyridis: The Influence of Prey Mobility and Prey Species

The influence of prey mobility and species on prey selection by the coccinellid Harmonia axyridis Pallas was determined under laboratory conditions for two prey species, Hyaliodes vitripennis (Say) and Tetranychus urticae Koch. Prey selection was influenced by prey mobility. In the presence of active prey, the coccinellid selected T. urticae while in presence of immobilized prey, H. vitripennis was preferred. Harmonia axyridis searching time was longer in the presence of active H. vitripennis than in the presence of active T. urticae. Moreover, the coccinellid capture rate was lower for active H. vitripennis caused by effective defensive mechanisms. Prey suitability was affected by prey mobility and species. Immobilized H. vitripennis were the most profitable prey, i.e. induced a shorter developmental time and no mortality. However, active H. vitripennis were not a suitable food source for H. axyridis. Our results suggested that three factors are involved in prey selection by H. axyridis: (i) prey mobility; (ii) prey defensive mechanisms; and (iii) prey species.     

Voracity and feeding preferences of two aphidophagous coccinellids on Aphis citricola and Tetranychus urticae

Voracity and feeding preferences of adult Coccinella septempunctata L. and Harmonia axyridis Pallas (Coleoptera: Coccinellidae, tribe Coccinellini) were evaluated in the laboratory on a common prey, the spirea aphid, Aphis citricola van der Goot (Homoptera: Aphididae), and on the twospotted spider mite, Tetranychus urticae Koch (Acarina: Tetranychidae), a prey previously unrecorded for these two predators. The experiments were carried out in the laboratory on apple saplings (Malus domestica Borkhausen). Adult males and females of H. axyridis consumed significantly more mites than adults of C. septempunctata. For H. axyridis, males consumed 41.3 spider mites in 24 h and females 48.4, whereas for C. septempunctata males consumed 14.1 prey and females 15.2. The consumption of spirea aphids by the males was similar for the two species. Consumption by the females was significantly greater for H. axyridis (46.5) than for C. septempunctata (22.2). The two coccinellids showed a significant preference for A. citricola in the presence of T. urticae. This preference was more pronounced for C. septempunctata. The total number of prey consumed and the percentage of exploited biomass decreased significantly for C. septempunctata and stayed relatively constant for H. axyridis as the number of mites increased in the prey ratio. Our results suggest that T. urticae is only an alternative prey for both predators, and that H. axyridis should be more efficient than C. septempunctata in a prey assemblage with aphids and mites.     

Odour-mediated preference and prey preference of Macrolophus caliginosus between spider mites and green peach aphids

Abstract  A comparative study between direct prey preference and odour-mediated preference of the predatory mirid bug, Macrolophus caliginosus , was conducted. We used a Y-tube olfactometer to determine the attraction of the predator to herbivore-induced volatiles from Myzus persicae or Tetranychus urticae -infested sweet pepper plants over clean plants and to direct prey odours over clean air. The mirid bugs showed a stronger response to odours from infested plants than to odours from clean plants. The mirids did, however, not seem to exploit odours emitted directly from the prey themselves. Our results further demonstrated that M. caliginosus prefers M. persicae to T. urticae in a direct two-choice consumption test. This preference was, however, not reflected in a similar odour-mediated preference between plant volatiles induced by either of the two preys. Two hypotheses are suggested for this neutral response of the mirids to herbivore-induced volatiles from a preference prey or a non-preference prey.     

Patterns, mechanisms and spatial scale of aggregation in generalist and specialist predatory mites (Acari: Phytoseiidae)

Prey species often distribute themselves patchily in their habitats. In response to this spatial variation in prey density, some predator species aggregate in patches of higher prey density. This paper reviews a series of laboratory experiments to demonstrate the patterns of responses by phytoseiid predators (Phytoseiulus persimilis, Typhlodromus occidentalis and Amblyseius andersoni) to spatial variation in the density of their spider mite prey (Tetranychus urticae) and reveal the behavioural mechanisms underlying the observed patterns. In addition, patterns of aggregation were examined at a variety of spatial scales on plants in greenhouses. The patterns, mechanisms and spatial scale of aggregation in three predatory species are discussed in relation to their varying degrees of polyphagy. The results show that a specialist predator species (1) aggregates more strongly than generalist predators, (2) does so not because it finds prey patches of high density more easily but because it remains in these patches longer than generalist predators and (3) tends to aggregate more often at lower levels of spatial scale than generalist predators. It is suggested that these conclusions, based mainly on laboratory studies of a small sample of species, should be tested in the future on a wider selection of specialist and generalist species at different scales in the field. This revised version was published online in November 2006 with corrections to the Cover Date.