Grape downy mildew Plasmopara viticola, an alternative food for generalist predatory mites occurring in vineyards

Generalist phytoseiids are often observed for long periods on plants in the absence of prey, feeding on alternative foods and reaching high population levels. The persistence of generalist predatory mites on plants with a scarcity or absence of prey is a requirement for successful biocontrol strategies of herbivore mites. The importance of pollen as an alternative food for the support of generalist predatory mite populations is widely recognized. However, on grape the presence of pollen is often limited and thus other food sources should contribute towards generalist predatory mite persistence on perennial plants. Previous field observations reported the relationships between the population increases of generalist phytoseiids with late-season spread of grape downy mildew (GDM) Plasmopara viticola. In this study, we test the hypothesis that GDM could be a suitable food source for the predatory mites Amblyseius andersoni and Typhlodromus pyri. In the laboratory we compared the development times, oviposition rates and life-table parameters of predatory mites feeding on pollen or GDM mycelium and spores. Grape downy mildew supported the survival, development and oviposition of T. pyri and A. andersoni. Life-table parameters showed that GDM was a less suitable food source than pollen for both phytoseiid species and that it was more favorable for A. andersoni than for T. pyri. Implications for predator–prey interactions and conservation biological control in vineyards are discussed.     

Biological control of broad mites (<Emphasis Type="Italic">Polyphagotarsonemus latus</Emphasis>) with the generalist predator <Emphasis Type="Italic">Amblyseius swirskii</Emphasis>

The broad mite is a serious pest of a variety of crops worldwide. Several phytoseiid mites have been described to control these mites. However, broad mites are still one of the major pest problems on greenhouse pepper in South-eastern Spain. The generalist predatory mite A. swirskii is widely used against other pests of pepper plants such as thrips and whiteflies, the latter being a vector of broad mites. We assessed the potential of A. swirskii to control broad mites. The oviposition rate of A. swirskii on a diet of broad mites was lower than on a diet of pollen, but higher than oviposition in the absence of food. Population-dynamical experiments with A. swirskii on single sweet pepper plants in a greenhouse compartment showed successful control of broad mites.     

Pollen as Food for the Predatory Mites Iphiseius Degenerans and Neoseiulus Cucumeris (Acari: Phytoseiidae): Dietary Range and Life History

Although all known phytoseiid mites (Acari: Phytoseiidae) are predators of mites or small insects, many readily feed and reproduce on pollen as well. This ability to feed on food from plant origin increases their survival during periods when prey is locally sparse, but might occur at the expense of the ability to utilize food as efficiently as specialized predators. In this study we compare two predatory mite species used as biological control agents against thrips, Neoseiulus cucumeris and Iphiseius degenerans, with respect to (1) the range of pollen species that may serve as food sources for a sustained oviposition; and (2) the life history and expected intrinsic growth rates on some suitable pollen diets. The results show that I. degenerans is, compared to N. cucumeris, able to utilize a larger proportion of approx. 25 pollen species tested, but does not show equally high ovipositional rates as N. cucumeris. Consequently, the highest intrinsic growth rate for I. degenerans (0.21 day ^–1) will be surpassed by N. cucumeris.     

Juvenile prey induce antipredator behaviour in adult predators

It is generally assumed that the choice of oviposition sites in arthropods is affected by the presence of food for the offspring on the one hand and by predation risk on the other hand. But where should females oviposit when the food itself poses a predation risk for their offspring? Here, we address this question by studying the oviposition behaviour of the predatory mite Amblyseius swirskii in reaction to the presence of its counterattacking prey, the western flower thrips Frankliniella occidentalis. We offered the mites a choice between two potential oviposition sites, one with and one without food. We used two types of food: thrips larvae, which are predators of eggs of predatory mite but are consumed by older predator stages, and pollen, a food source that poses no risk to the predators. With pollen as food, the predators preferred ovipositing on the site with food. This might facilitate the foraging for food by the immature offspring that will emerge from the eggs. With thrips as food, female predators preferred ovipositing on the site without thrips. Predators that oviposited more on the site with thrips larvae killed more thrips larvae than females that oviposited on the site without food, but this did not result in higher oviposition. This suggests that the females killed thrips to protect their offspring. Our results show that predators display complex anti-predator behaviour in response to the presence of counter-attacking prey.     

Ecological and genetic factors influencing evolution of pesticide resistance in tetranychid and phytoseiid mites

Among tetranychid spider mites and their phytoseiid predators, the evolution of pesticide resistance is a common event. In most cases, resistance is based on a single dominant or semidominant gene. However, polygenic, less-stable resistance often develops under laboratory selection. More rapid development of pesticide resistance in spider mites and predatory mites than among other arthropods might partly be due to their arrhenotokous reproduction. For both groups of mites, little study has been done on population genetic factors influencing pesticide resistance. A few studies have focussed on ecological factors. An important ecological factor influencing resistance evolution is the level of immigration of susceptible individuals into treated habitats. Spider mites and predatory mites both tend to reside in treated habitats at high levels and to immigrate at only modest levels from untreated habitats. This favors rapid resistance development. Another factor contributing to rapid resistance evolution in both mite groups is their rapid reproductive rate. A food-limitation factor may limit resistance evolution under field conditions more in predatory mites than spider mites. After treatment by a pesticide, spider mites have an unlimited food source, whereas predatory mites have a decimated food source (their prey), which leads to reduced reproduction, starvation, or migration. Because of the common occurrence of resistance among both mite groups, a strategy of resistance management is often feasible for them. Case histories of IPM where the population dynamics and genetics of pesticide resistance of tetranychid and phytoseiid mites have been considered are discussed. The overall conclusion is that greater understanding of the population genetics and ecology of these species will provide for improved systems of resistance management and IPM.     

Provision of astigmatid mites as supplementary food increases the density of the predatory mite Amblyseius swirskii in greenhouse crops,but does not support the omnivorous pest,western flower thrips

Astigmatid mites can be used as prey for mass rearing of phytoseiid predators, but also as a supplemental food source to support predator populations in crops. Here we evaluated the potential of six species of astigmatid mites (living or frozen) as alternative food for the predatory mite Amblyseius swirskii Athias-Henriot in greenhouse crops. All prey mites tested were suitable for predator oviposition. In general, oviposition was greater when prey mites were reared on dog food with yeast than when they were reared on wheat bran with yeast. Amongst prey items provided as frozen diet, larvae of Thyreophagus entomophagus (Laboulbene), Acarus siro L. and Lepidoglyphus destructor (Schrank) that had been reared on dog food with yeast, resulted in the highest oviposition rates of A. swirskii. T. entomophagus larvae as frozen diet resulted in the shortest preimaginal developmental time of A. swirskii. On chrysanthemum plants, we found that the greatest increase in predator density occurred when living mites of T. entomophagous were used as a food source. This increase was greater than when predators were fed cattail pollen, a commonly used supplemental food. Effects on predators of providing living A. siro and L. destructor, or frozen larvae of T. entomophagous as food, were comparable with provision of pollen. Use of supplemental food in crops can be a risk if it is also consumed by omnivorous pests such as western flower thrips, Frankliniella occidentalis Pergande. However, we showed that both frozen and living mites of T. entomophagous were unsuitable for thrips oviposition. Hence, we believe that provision of prey mite species increases A. swirskii density, supporting biological control of thrips and other pests in greenhouse crops.

     

Predatory mites avoid ovipositing near counterattacking prey

Attacking prey is not without risk; predators may endure counterattackby the prey. Here, we study the oviposition behaviour of a predatory mite(Iphiseius degenerans) in relation to its prey, thewesternflower thrips (Frankliniella occidentalis). This thrips iscapable of killing the eggs of the predator. Thrips and predatory mites - apartfrom feeding on each other - can also feed and reproduce on a diet of pollen.Because thrips may aggregate at pollen patches, such patches may be risky foroviposition by the predatory mites. We found that, in absence of thrips,predatory mites lay their eggs close to pollen, but further away when thripsarepresent. Predatory mite eggs near pollen were killed more frequently by thripsthan when they were deposited further away. The oviposition behaviour of thepredatory mite was also studied in absence of thrips, but in presence of thealarm pheromone of thrips. This pheromone is normally secreted upon contactwithpredators or competitors. When applied close to the pollen, predatory mitesoviposited significantly further away from it. When the alarm pheromone wasapplied away from the food source, most eggs were found near the pollen. Theseresults indicate that female predatory mites show flexible ovipositionbehaviourin response to the presence of their counterattacking prey.     

Abundance of phytoseiid mites on Vitis species: effects of leaf hairs,domatia, prey abundance and plant phylogeny

We observed the number of predatory mites (Phytoseiidae:Typhlodromus caudiglans) on the foliage of 20 North American species of grapes (Vitis spp) plus the domesticated EuropeanVitis vinifera, all grown in a common garden. We found relatively few phytophagous mites. The numbers of phytophagous mites were not correlated with the plant characteristics that we measured. We found approximately five times as many predatory mites as phytophagous mites and the numbers of these phytoseiid predators were not affected by the availability of prey. Similarly, numbers of phytoseiids were unaffected by plant gender and, hence, the availability of pollen, another source of food. The numbers of phytoseiids were not clustered according to the taxonomic grouping of the tested plant species. Leaf surface characteristics explained over 25% of the variance in the numbers of phytoseiids. Numbers of phytoseiids were positively associated with the density of vein hairs, the density of bristles in leaf axils, and the presence of leaf domatia. These results suggest that sheltered habitats rather than food availability may limit the numbers of phytoseiid mites on grapevines.     

A laboratory study on the predatory miteTyphlodromus pyri (Acarina: Phytoseiidae): I. The effect of temperature and food consumption on the rate of development of the eggs and immature stages

Development periods were determined for eggs and juvenile stages of the predatory phytoseiid mite Typhlodromus pyri at four constant temperatures and three feeding levels. For the non-feeding stages (eggs and larvae) power curve functions were fitted to development rate in terms of temperature. The results are compared to the general arthropod response to temperature. A non-linear model was fitted to nymphal development rate in terns of temperature and food, in which the weight gain over an instar is assumed to be a linear function of daily food consumption. Examples of the same shape development rate—food relationship are given for two more phytoseiid mites and for other arthropods. The results are discussed with respect to T. pyri as an integrated control agent.     

Grape powdery mildew as a food source for generalist predatory mites occurring in vineyards: effects on life-history traits

In several perennial cropping systems, generalist or omnivorous species represent important biocontrol agents. They can persist on plants by feeding on alternative foods when prey is scarce and potentially limit pest outbreaks. Among beneficials characterised by a wide food range, those belonging to the acarine family Phytoseiidae represent important biocontrol agents. Generalist predatory mites can develop and reproduce using various food sources as alternatives to their tetranychid prey. The presence of alternative food sources can also induce switching feeding behaviour of generalist predators from prey to alternative foods. We evaluated in the laboratory the role of the grape powdery mildew (GPM) for the survival, development and reproduction of Amblyseius andersoni and Typhlodromus pyri , two important beneficial phytoseiid mites, in European and North-American vineyards. We also compared life-history parameters obtained when feeding on GPM with those obtained feeding on tetranychids mite prey or cattail pollen. Results indicated that GPM is an adequate food source for generalist mite survival and development. Results suggest that GPM can sustain mite populations in the absence of higher quality food sources. Based on optimal foraging theory, comparison of life-history parameters on GPM and mite prey suggests that the disruption of phytophagous mite control by these predatory mites in the presence of GPM appears unlikely. Implications for biological control in vineyards are discussed.     

Herbivory-associated degradation of tomato trichomes and its impact on biological control of Aculops lycopersici

Tomato plants have their leaves, petioles and stems covered with glandular trichomes that protect the plant against two-spotted spider mites and many other herbivorous arthropods, but also hinder searching by phytoseiid mites and other natural enemies of these herbivores. This trichome cover creates competitor-free and enemy-free space for the tomato russet mite (TRM) Aculops lycopersici (Acari: Eriophyidae), being so minute that it can seek refuge and feed inbetween the glandular trichomes on tomato cultivars currently used in practice. Indeed, several species of predatory mites tested for biological control of TRM have been reported to feed and reproduce when offered TRM as prey in laboratory experiments, yet in practice these predator species appeared to be unable to prevent TRM outbreaks. Using the phytoseiid mite, Amblydromalus limonicus, we found exactly the same, but also obtained evidence for successful establishment of a population of this predatory mite on whole plants that had been previously infested with TRM. This successful establishment may be explained by our observation that the defensive barrier of glandular plant trichomes is literally dropped some time after TRM infestation of the tomato plants: the glandular trichome heads first rapidly develop a brownish discoloration after which they dry out and fall over onto the plant surface. Wherever TRM triggered this response, predatory mites were able to successfully establish a population. Nevertheless, biological control was still unsuccessful because trichome deterioration in TRM-infested areas takes a couple of days to take effect and because it is not a systemic response in the plant, thereby enabling TRM to seek temporary refuge from predation in pest-free trichome-dense areas which continue to be formed while the plant grows. We formulate a hypothesis unifying these observations into one framework with an explicit set of assumptions and predictions to be tested in future experiments.     

Pollen on-twine for food provisioning and oviposition of predatory mites in protected crops

The establishment of predatory mites on protected crops is affected by the availability of shelter and alternative food. We suggest that a pollen covered twine, coined here “pollen on-twine”, may provide these necessities when attached to the plant. We evaluated the effect of twine types and two pollen species on the establishment of Euseius scutalis (Athias-Henriot) and Amblyseius swirskii Athias-Henriot on pre-flowering pepper plants in a growth chamber. For both phytoseiids, rayon jute twine was found more beneficial while corn and oak pollen did not differ in their effect. Though populations of both predators were best promoted when twine and pollen were applied separately, E. scutalis population increased by more than tenfold and A. swirskii doubled when plants were applied with pollen on-twine. We propose that after further refining, pollen on-twine can serve as a feasible solution for predatory mite establishment on protected crops.     

中国植绥螨规模化饲养及保护利用研究进展

根据植绥螨食性的不同 ,可以采用叶螨、花粉和人工饲料等食物来培养植绥螨。规模化饲养植绥螨多用叶螨等害螨的寄主植物先繁殖害螨 ,然后在害螨中加入一些补充食物来大量生产植绥螨。张艳璇设计的袋栽法就是这样一套新的大量繁殖植绥螨的工艺方法。植绥螨的贮存目前都采用降低发育来实现 ,一般在 5～ 1 0℃的温度下能保存 5 0d以上。在田间按一定的比例释放抗药性的植绥螨 ,能够有效的控制二斑叶螨TetranychusurticaeKoch等害螨的发生。并且在地面鼓励种植覆盖作物、允许少量杂草生长或实行生草栽培 ,能够为植绥螨提供交替食物 ,从而更好地发挥生态控制功能的作用。     

Conspicuous extra-floral nectaries are inducible in Vicia faba

Mutualistic interactions are dynamic associations that vary depending on the costs and benefits to each of the interacting parties. Phenotypic plasticity in mutualistic interactions allows organisms to produce rewards to attract mutualists when the benefits of their presence outweigh the costs of producing the rewards. In ant–plant defensive mutualisms, defences are indirect as plants produce extra‐floral nectaries (EFN) to attract predatory ants to deter herbivores. Here we demonstrate that in broad bean, Vicia faba, the overall number of EFNs on a plant increases dramatically following leaf damage. In two damage treatments, removal of: (1) one‐third of one leaf in a single leaf pair or (2) one‐third of both halves of a single leaf pair, resulted in a 59 and 106% increase in the number of EFNs on the plants, respectively, over 1 week. We suggest that the increased production of visually conspicuous EFNs is an adaptive inducible response, to attract predatory arthropods when risk of herbivory increases.     

Assessing the Effects of Bt Maize on the Predatory Mite Neoseiulus cucumeris

The investigation of Neoseiulus cucumeris in the context of the ecological risk assessment of insect resistant transgenic plants is of particular interest as this omnivorous predatory mite species is commercially available and considered important for biological control. In a multitrophic feeding experiment we assessed the impact of Bt maize on the performance of N. cucumeris when offered spider mites (Tetranychus urticae) reared on Bt (Bt11, Syngenta) or non-Bt maize (near isogenic line) and Bt or non-Bt maize pollen as a food source. Various parameters including mortality, development time, oviposition rate were measured. Spider mites were used as a prey for N. cucumeris, since these herbivores are known to contain similar levels of Cry1Ab toxin, when reared on Bt maize, as those found in the transgenic leaf material. In contrast, toxin levels in pollen of this transgenic cultivar are very low. No differences in any of the parameters were found when N. cucumeris was fed with spider mites reared on Bt and non-Bt maize. Pollen was shown to be a less suitable food source for this predator as compared to spider mites. Moreover, subtle effects on female N. cucumeris (9% longer development time and 17% reduced fecundity) were measured when fed with pollen originating from Bt maize as compared to non-Bt maize pollen. Our findings indicate that the predatory mite N. cucumeris is not sensitive to the Cry1Ab toxin as no effects could be detected when offered Bt-containing spider mites, and that the effects found when fed with Bt maize pollen can be assigned to differences in nutritional quality of Bt and non-Bt maize pollen. The significance of these findings is discussed with regard to the ecological relevance for risk assessment of transgenic plants.     

中国植绥螨资源及其生物学研究进展

植绥螨是许多害螨和小型有害昆虫的重要捕食性天敌 ,在农业生产中极具利用价值。我国植绥螨资源相当丰富 ,大量的资源调查工作是在 2 0世纪 70年代末开始的。到目前为止 ,我国已经从不同地区作物上鉴定出植绥螨 2 60余种 ,其中有利用价值的约 2 0种左右。在 2 5～ 3 0℃的适宜温度下 ,植绥螨从卵到羽化为成螨约需 6～ 7d ,而寿命能长达 3 0～ 5 0d。它们对大多数的害螨的功能反应基本成HollingⅡ型。多数成螨为喜湿类型 ,较高的湿度对于孵化和存活尤为重要。在夏季高温和冬季低温来临时进行越夏和越冬 ,以度过不良的环境。植绥螨行两性生殖 ,不交配或交配时间不够 ,不产卵或少产 ,有些种类需要多次交配才能完全产卵。     

Effect of different plant pollens on the development and oviposition of seven native phytoseiid species (Acari: Phytoseiidae) in Japan

The quality of pollen of tea (Camellia sinensis L.), big leaf podocarp tree [Podocarpus macrophyllus (Thumb.)] and sweet corn (Zea mays L.) as alternative food sources was evaluated for seven native phytoseiid mite species, Amblyseius eharai Amitai and Swirski, Euseius sojaensis (Ehara), Neoseiulus californicus (McGregor), Neoseiulus womersleyi (Schicha), Phytoseius nipponicus Ehara, Chanteius contiguus (Chant) and Typhlodromus vulgaris Ehara. Tea pollen was of high nutritional value for all seven phytoseiid species: most larvae developed to adults and most females oviposited well. Development and oviposition differed greatly among phytoseiid species when reared on big leaf podocarp tree pollen: A. eharai, T. vulgaris and C. contiguus could utilize the pollen as food, but the other phytoseiid species could not. In addition, big leaf podocarp tree pollen could be utilized by A. eharai as food for longer than the other pollen. The quality of sweet corn pollen was inferior in general, although it could somehow favor the development and oviposition of six phytoseiid species except for E. sojaensis.     

The Contribution of Extrafloral Nectar to Survival and Reproduction of the Predatory Mite Iphiseius Degenerans on Ricinus Communis

The phytoseiid mite Iphiseius degenerans (Berlese) is an effective predator of western flower thrips, Frankliniella occidentalis (Pergande), in Dutch greenhouses. In the Mediterranean area, castor bean, Ricinus communis L., is known as a year-round host plant for this predatory mite. On flowering castor bean plants in greenhouses, I. degenerans can be found in densities of more than 100 per leaf. For this reason, the plant is being used as a banker plant to augment biological control. It has been shown that pollen produced by the large apical flowers sustains reproduction and development for these mites. The objective of this study was to measure the contribution of the extrafloral nectar of this plant to the reproductive success of this predatory mite. A study conducted at 25°C in presence of free water showed that (1) I. degenerans is unable to develop beyond the protonymphal stage when fed only nectar and leaf tissue, (2) its ovipositional rate is higher when pollen is supplemented with nectar, (3) its reproduction ceases within a few days when fed on nectar only, but the predator can survive for several weeks and resume oviposition when fed pollen again and (4) the feeding of young females for one or two weeks with nectar only extends their longevity by approximately the same period and only slightly diminishes their lifetime reproductive potential (R0), as compared to mites continuously fed pollen. It can be concluded that extrafloral nectar can provide an important contribution to population growth and maintenance of I. degenerans on R. communis, particularly in pre- and post-blooming periods. Assuming these predators are beneficial to the plant in clearing them of herbivorous mites and thrips, this relationship may be regarded as an example of plant–predator mutualism. The combination of pollen and extrafloral nectar makes castor bean an ideal rearing and banker plant for I. degenerans.     

Artificial ground shelters for overwintering phytoseiid mites in orchards

Biological control in orchards strongly depends on winter survival of natural enemies, especially in temperate regions. Predacious phytoseiid mites overwinter on trees or on the ground depending on the characteristics of the species. However, the overwintering ecology of phytoseiid mites on the ground is less well known than that of those on trees. We investigated the usefulness of artificial overwintering shelters as a tool for studying the overwintering ecology of phytoseiid mites on the ground. Four kinds of artificial shelter (shading net, felt, cardboard, and urethane foam) were placed on the ground in an apple orchard in Korea. Two dominant phytoseiid species, Neoseiulus makuwa (Ehara) and N. womersleyi (Schicha) (Acari: Phytoseiidae), overwintered in the artificial ground shelters, and numbers were highest in the urethane foam among the four kinds of shelter, and next highest in the shading net. On the other hand, the numbers of phytoseiid mites collected in the ground vegetation plus soil samples under the ground shelters were not significantly different among the five shelter treatments, including the no-shelter control. Our results suggest that artificial overwintering shelters are efficient tools for investigating overwintering ecology of phytoseiid mites on the ground, as well as on trees, in orchards. Furthermore, the artificial shelters would be good sampling units because they are easily formed into identical sizes and can be used almost anywhere in the field with less laborious work. We also discuss some implications about the effects of sheltered structures on the ground on the populations of phytoseiid mites during winter.     

Phytophagous and predatory mites (Acari: Tetranychidae,Eriophyidae, Phytoseiidae,Stigmaeidae) in South Moravian vineyards,Czechoslovakia, treated with various types of chemicals

Communities of phytophagous and predatory mites on vine can be influenced by the type of chemical treatment. Ten species of phytoseiid mites inhabit vines in the region of South Moravia. Populations ofTyphlodromus pyri Scheuten play leading roles in effective suppression of tetranychid and eriophyid mites in commercial vineyards sprayed with pesticides, except synthetic pyrethroids and mancozed, which are considered to be detrimental to the predatory phytoseiid mites.