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.     

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.     

Mite Population Dynamics on Different Grape Varieties With or Without Phytoseiids Released (Acari: Phytoseiidae)

In a three-year study, mite populations were monitored in two vineyards, each having two grape varieties with different leaf hair density. In both vineyards native phytoseiids were present: Amblyseius andersoni in one vineyard, and Phytoseius finitimus in the other. The economically important predators Kampimodromus aberrans and Typhlodromus pyri were released in both vineyards in order to study their efficacy in controlling tetranychids and eriophyids and their persistence during periods of prey scarcity. In both vineyards, relative abundances of the mite species, especially phytoseiids, were found to differ on different varieties in the same vineyard. In the first experiment, A. andersoni reached higher densities and was more persistent on the variety with slightly pubescent leaf under-surface (Merlot). Typhlodromus pyri and K. aberrans releases were successful and the mites became more abundant on the variety with pubescent leaf under-surface (Verduzzo). In the second experiment, P. finitimus was more abundant on a variety with pubescent leaf under-surface (Prosecco) than with glabrous leaf under-surface (Riesling). The most interesting results of the present study concerned the interactions between native and released predators. In the first vineyard, different results were obtained when releasing T. pyri on the two varieties. On the variety with pubescent leaves, A. andersoni was rapidly displaced by T. pyri, whereas the former species persisted on the other variety throughout the three-year study, apparently becoming dominant during the last season. In contrast to T. pyri, interactions between K. aberrans and A. andersoni in this vineyard did not depend on variety. The results of the experiments carried out in the second vineyard stressed the importance of interspecific competition for phytoseiid releases. Typhlodromus pyri colonization failed on both varieties. Kampimodromus aberrans releases appeared to be more successful on Riesling than on Prosecco, where P. finitimus was more abundant. At the end of the experiments, K. aberrans displaced P. finitimus on both varieties.     

Behavioral and population consequences of acarodomatia in grapes on phytoseiid mites (Mesostigmata) and implications for plant breeding

We examined the influence of acarodomatia in the riverbank grape Vitis riparia Michaux (Vitaceae) on the distribution and abundance of predatory mites (Phytoseiidae) and their interactions with herbivorous mites. Acarodomatia are tufts of nonglandular trichomes or pits located in major leaf vein axes of many species of woody perennial plants and are often occupied by predatory and mycophagous mites. In common garden plantings of different accessions of V. riparia we found a significant positive relationship between size of domatia and the abundance of naturally occurring predatory mites. Behavior of adult predatory mites may explain this positive association, in part. In separate laboratory experiments, gravid females of Typhlodromus pyri Scheuten and Amblyseius andersoni Chant spent more time and deposited more eggs on half of a V. riparia leaf with accessible domatia versus the other half in which access to domatia was blocked with pruning tar. Domatia also had population consequences. In an outdoor experiment using potted grapevines, population size of T. pyri and A. andersoni mites was greater on V. riparia with open domatia compared to V. riparia in which domatia were blocked with pruning tar. Population size of predatory mites was also greater on V. riparia with domatia than on Vitis vinifera L., whether their axils were blocked or not. Since V. vinifera have very small domatia, these results indicate that the presence of domatia is important, not just access to vein axils. Elevated predatory mite populations in response to domatia, however, did not translate into differences in the abundance of European red mite Panonychus ulmi (Koch), an important pest of grapes.Overall, these results indicate that domatia in uncultivated V. riparia promote higher densities of some species of generalist phytoseiid mites. However, domatia are small in most cultivated grapes. We crossed females and males of V. riparia that varied in domatia size and reared their offspring and found that average domatia size in the parents was highly correlated (r² = 0.77, slope = 0.55) with average domatia size in offspring (high narrow-sense heritability). Given that V. riparia possesses many other desirable agronomic traits, this result suggests it should be practical to breed for well-developed domatia in cultivated accessions.     

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.     

Plants,mites and mutualism: leaf domatia and the abundance and reproduction of mites on Viburnum tinus (Caprifoliaceae)

Associations between mites and leaf domatia have been widely reported, but little is known about their consequences for either plants or mites. By excising domatia from leaves of the laureltinus, Viburnum tinus L. (Caprifoliaceae), in the garden and laboratory, we showed that domatia alter the abundance, distribution, and reproduction of potential plant mutualists. Over 4 months, leaves with domatia on six garden shrubs had 2–36 times more predatory and microbivorous mites, and more mite eggs than leaves without domatia. However, this effect varied among plants and was weaker on one shrub with few mites on its leaves. Domatia also influenced the distribution of mites on leaves. A significantly higher fraction of mites, representing all life stages, was found in vein axils of leaves with domatia than in vein axils on leaves without domatia. Single-leaf experiments in the laboratory showed that domatia enhanced reproduction by the predatory mite, Metaseiulus occidentalis, especially at low relative humidity (30–38%). When domatia were removed, oviposition was reduced significantly only at low relative humidity, suggesting that domatia provide mites with refuge from environmental extremes on the leaf surface. Moreover, the use of domatia by predatory mites may reduce the impact of some plant enemies. In two experiments where prey consumption was measured, M. occidentalis ate significantly higher percentages of the eggs of the two-spotted spider mite (Tetranychus urticae). Our results are consistent with the viewpoint that mite-domatia associations are mutualistic. By directly aiding and abetting the third trophic level, plants with leaf domatia may increase the efficiency of some predaceous and microbivorous mites in consuming plant enemies.     

Colonization of Languedoc vineyards by phytoseiid mites (Acari: Phytoseiidae): influence of wind and crop environment

A natural increase of phytoseiid mite populations (Kampimodromus aberrans, Typhlodromus pyri and Phytoseius plumifer) was observed in vineyards in Languedoc, Burgundy and Corsica under integrated pest management strategies. The aim of the present study was to characterize the mechanisms of this colonization in space and time in Languedoc. The abundance of phytoseiid mites in the vegetation close to three grape fields was determined twice a year (May and July). Aerial (funnels with water) and soil (felt strip) traps were placed in and around grape fields, in order to assess the colonization potential provided by aerial dispersal and ambulatory locomotion. The populations of phytoseiid mites in the crops were studied twice a month in order to gain information on the make up of the dispersal populations. The species K. aberrans was found in largest quantities in the traps, in the natural vegetation and in the crops. Predatory mite dispersal occurred essentially by aerial dispersal and was dependent on the wind intensity and wind direction. Identical sex ratios were observed in migrant populations and in populations present in the grape fields, woody areas and hedges. A large proportion of immatures was found to move by aerial dispersal. The colonization potential (rapidity, intensity and regularity) was directly associated with the abundance of the phytoseiids and the proximity of natural vegetation. A deep, dense and tall woody area containing suitable host plants for predatory mites constituted the most stable source of phytoseiid mites. Natural colonization of vineyards provides considerable phytoseiid mite potential that could be managed in an agricultural landscape.     

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.     

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.     

Leaf trichomes influence predatory mite densities through dispersal behavior

Habitat complexity can mediate interactions among predators and herbivores and influences arthropod population density and community structure. The abundance of many predatory mites (Acari: Phytoseiidae) is positively associated with abundance of non‐glandular trichomes. We hypothesized that (1) increasing the complexity (trichome density mimicked with cotton fiber patches) of the habitat that predatory mites encounter on leaves would reduce adult dispersal from plants, and (2) increasing habitat complexity would reduce the time that mites spend walking. Typhlodromus pyri Scheuten retention on plants increased linearly in the presence of trichome mimics; mites placed on plants lacking leaf trichomes showed a behavioral response that led to active dispersal. Phytoseiid retention increased with both fiber patch size and fiber density within patches. Moving fiber patches from the underside of the leaf to the upper leaf surface did not change phytoseiid retention but did alter egg distribution, suggesting trichomes do not exclusively influence phytoseiid behavior. Phytoseiid activity level as measured by the amount of time spent walking did not decrease with the addition of fibers. Overall, increasing habitat complexity in the form of non‐glandular trichomes strongly reduced T. pyri dispersal behavior; the predatory mites showed a consistent preference for complex trichome‐rich habitat that was manifest both rapidly and in absence of predators. Hence, the frequently observed pattern of population‐level accumulation of phytoseiids on trichome‐rich plants appears to be driven by a behavioral response to the presence and abundance of non‐glandular trichomes on the leaf surface manifested in the level of dispersal and/or retention. The primary implication of phytoseiid–habitat interactions for biocontrol programs is that where plants have no trichomes, T. pyri will not establish. Whether this behavioral response pattern is a general response of phytoseiids to leaf trichomes or varies with species is a question that remains unanswered.     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

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.     

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.     

Leaf domatia and mites on Australasian plants: ecological and evolutionary implications

Leaf domatia, specialized chambers in the vein axils on the underside of leaves of many plant species, have remained an enigma for over a century. In this study we show a strong association between foliar domatia and mites in 37 plant species in Australasia. Overall, mites accounted for 91% of the arthropods observed in domatia. Across all species, a median of 51% of domatia were occupied and 71% of leaves showed mite evidence in domatia. The level of mite association did not depend on domatia type (pit, pouch, pocket, or tuft) or provenance (Papua New Guinea, Queensland, Victoria, or New Zealand). Mite association with domatia commonly varied between plant species, between individuals within species, and between shoots within individuals. The leaf developmental stage probably explains much of the variation in association for many of these species. The presence of a variety of life history stages of mites within domatia indicates that these structures act as shelters for development and reproduction. Furthermore, in 12 of 13 plant species examined, domatia concentrate mites in particular locations on the leaf. Mite taxa that we classify as largely predaceous (e.g. phytoseiids, stigmaeids and tydeids) or fungivorous (e.g. acarids and oribatids) were most common in domatia and dominated the association in 21 of 24 plant species in which the relative abundance of herbivorous, fungivorous and predaceous groups was quantified. We evaluate hypotheses that explain the role of leaf domatia, including non-functional hypotheses (e.g. architectural constraints), physiological function (e.g. gas exchange and water uptake), bacterial symbiosis and antagonistic and mutualistic associations with mites. Our quantitative results confirm anecdotal accounts of mite association with leaf domatia and are most consistent with Lundströem's century-old hypothesis of plant-mite mutualism in which leaf domatia billet predaceous and fungivorous mites that prey on plant enemies. Leaf domatia are widespread among woody angiosperms and abundant in many temperate and tropical regions of Australasia. Mites, an ancient group of arthropods whose diversity and abundance parallels that of insects, are likely to be important selective agents on terrestrial plants. Our results (1) indicate that mite-domatia association represents a relationship of comparable scope to plant-ant associations mediated by specialized plant structures such as extrafloral nectaries, food bodies and specialized domatia; (2) suggest that sociality is not a necessary prerequisite for widespread and diverse mutualisms between arthropods and plants; and, (3) extend the diversity of organisms that produce specialized mite ‘houses’ from lizards, and wasps and bees to woody angiosperms.     

Leaf pubescence mediates the abundance of non-prey food and the density of the predatory mite Typhlodromus pyri

Plants with leaves having numerous trichomes or domatia frequently harbor greater numbers of phytoseiid mites than do plant with leaves that lack these structures. We tested the hypothesis that this pattern occurs, in part, with Typhlodromus pyri because trichomes increase the capture of pollen or fungal spores that serve as alternative food. Using a common garden orchard, we found that apple varieties with trichome-rich leaves had 2–3 times more pollen and fungal spores compared to varieties with trichome-sparse leaves. We also studied the effects of leaf trichome density and pollen augmentation on T. pyri abundance to test the hypothesis that leaf trichomes mediate pollen and fungal spore capture and retention and thereby influence phytoseiid numbers. Cattail pollen (Typha sp.) was applied weekly to mature ‘McIntosh’ and ‘Red Delicious’ trees grown in an orchard and, in a separate experiment, to potted trees of the same varieties. ‘McIntosh’ trees have leaves with many trichomes whereas leaves on the ‘Red Delicious’ trees have roughly half as many trichomes. With both field-grown and potted trees, adding cattail pollen to ‘Red Delicious’ trees increased T. pyri numbers compared to ‘Red Delicious’ trees without pollen augmentation. In contrast, cattail pollen augmentation had no effect on T. pyri populations on ‘McIntosh’ trees. Augmentation with cattail pollen most likely supplemented a lower supply of naturally available alternative food on ‘Red Delicous’ leaves and thereby enhanced predator abundance. These studies indicate that larger populations of T. pyri on pubescent plants are due, in part, to the increased capture and retention of pollen and fungal spores that serve as alternative foods. 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.     

OCCURRENCE OF PREDATORY AND FUNGIVOROUS MITES IN LEAF DOMATIA

Domatia are small invaginations and hair tufts usually found at vein junctions on the undersides of leaves in many woody dicots. Domatia of 32 plant species (of worldwide origin from 18 families) growing in California, Hawaii, and Costa Rica were examined for mites. Domatia of 31 of 32 (97%) of these plants contained mites, and 24 of 32 (75%) contained mite eggs. Mites were found within the domatia of 48% of the sampled leaves. The domatia of 26 of 31 (84%) plants had mite species considered beneficial (primarily in the families Phytoseiidae and Tydeidae, but also Bdellidae, Cheyletidae and Stigmaeidae), while 6 of 31 (19%) had mite species considered harmful (Tenuipalpidae and Eriophyidae). Based on these findings (and in part upon Lundström's 1887 domatia theory), we hypothesize the existence of a widespread facultative mutualism between plants with leaf domatia and beneficial mites: leaf domatia serve as shelters and nurseries for mites which in turn eat phytophagous arthropods and pathogens using the plants. This proposed mutualism could be of importance to agriculture since domatia are known to occur in some crop plants, including coffee, grape, and walnut.     

Host plant manipulation of natural enemies: leaf domatia protect beneficial mites from insect predators

Acarodomatia are small tufts of hair or invaginations in the leaf surface and are frequently inhabited by several taxa of non-plant-feeding mites. For many years, ecologists have hypothesized that these structures represent a mutualistic association between mites and plants where the mites benefit the plant by reducing densities of phytophagous arthropods and epiphytic microorganisms, and domatia benefit the mite by providing protection from stressful environmental conditions, other predaceous arthropods, or both. We tested these hypothesized benefits of domatia to domatia-inhabiting mites in laboratory and growth chamber experiments. In separate experiments we examined whether domatia on the wild grape, Vitis riparia, provided protection against drying humidity conditions or predaceous arthropods to two species of beneficial mite: the mycophagous species Orthotydeus lambi, and the predaceous species Amblyseius andersoni. For both taxa of beneficial mite, domatia significantly increased mite survivorship in the presence of the predatory bug, Orius insidiosus and the coccinellids Coccinella septempunctata and Harmonia varigata. There was no evidence for a protective effect of domatia with a third species of predatory arthropod, lacewing larvae Chrysoperla rufilabris. In contrast, there was no evidence for either species of beneficial mite that domatia provided any protection against low humidity. Thus in this system the primary mechanism by which domatia benefit beneficial mites is by protecting these organisms from other predatory arthropods on the leaf surface.     

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.     

Resistance of grapevine to the erineum strain of Colomerus vitis (Acari: Eriophyidae) in western Iran and its correlation with plant features

The interaction of grape erineum mite (GEM), Colomerus vitis Pagenstecher (Acari: Eriophyidae), with grape was investigated in the laboratory. We studied some plant morphological biochemical features potentially related to vine resistance/tolerance of eight native grapevine cultivars, extensively cultivated in western Iran, and two non-native cultivars. Free-choice experiments indicated that the cultivars Shahani, Flame seedless and Yaghuti were colonized by lower levels of GEM, whereas Muscat Gordo, Gazne and White Thompson seedless hosted denser populations. These differences between cultivars may be due to differential attractiveness to GEM, possibly associated with plant biochemical and morphological traits. In no-choice assays with six grapevine cultivars, mite population development and some cultivar features were assessed. Mite populations grew fastest on Gazne and Muscat Gordo, and slowest on Yaghuti and Shahani. The degree of mite infestation was associated with reduction of leaf area, increase of leaf weight, shortening of shoots and more numerous erinea: these features were larger on the most infested Gazne, whereas morphological features of Shahani and Yaghuti were scarcely affected by GEM infestation. Also trichome type and density of the assayed cultivars appeared to be related to mite density: the most infested cultivars (Gazne and Muscat Gordo) displayed higher ranks of blade and vein hairs and lower ranks of blade and vein bristles and domatia. No correlation was found between mite density and leaf thickness of mature leaves. The amount of leaf waxes was highest in Shahani and Yaghuti, which displayed the lowest mite density, the fewest erinea and the largest leaves. Carbohydrate amount of uninfested leaves was lowest on the least infested Shahani and highest on the most infested Gazne; phenols increased in leaves of Shahani and decreased in those of Gazne after mite infestation. Finally, cultivars also appeared to influence some morphological traits of the mites: larger specimens were detected on White Thompson seedless, Flame seedless and Gazne, whereas smaller mites were found on leaves of the less infested Yaghuti and Shahani. These results indicate that leaf hairiness, leaf wax and carbohydrate contents may be useful tools for a preliminary screening among vine cultivars and help predict resistance/tolerance to GEM. Shahani and Yaghuti seem quite promising for developing grape resistance programs against GEM in western Iran.