Differential Effects of Plant Species on a Mite Pest (Tetranychus Urticae) and its Predator (Phytoseiulus Persimilis): Implications for Biological Control

The influence of plant species on the population dynamics of the spider mite pest, Tetranychus urticae, and its predator, Phytoseiulus persimilis, was examined as a prerequisite to effective biological control on ornamental nursery stock. Experiments have been done to investigate how the development, fecundity and movement of T. urticae, and the movement of P. persimilis were affected by plant species. A novel experimental method, which incorporates plant structure, was used to investigate the functional response of P. persimilis. Development times for T. urticae were consistent with published data and did not differ with plant species in a biologically meaningful way. Plant species was shown to have a major influence on fecundity (P < 0.001) and movement of the pest mite (P < 0.01), but no influence on the movement of the predator. The movement of both pest and predator was shown to be related to the density of the adult pest mites on the plant (P < 0.001). Plant structure affected the functional response, particularly in relation to the ability of the predator to locate prey at low densities. The impact of these findings on the effective use of biological control on ornamental nursery stock is 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.     

Evaluation of the predatory mite, Neoseiulus californicus, for spider mite control on greenhouse sweet pepper under hot arid field conditions

The efficacy of Neoseiulus californicus (a generalist predatory mite) for the biological control of Tetranychus urticae, was compared to release of Phytoseiulus persimilis (a specialist predatory mite) and an acaricide treatment in sweet pepper plants grown in greenhouse tunnels in a hot and arid climate. To ensure uniform pest populations, spider mites were spread on pepper plants in two seasons; a natural infestation occurred in one season. Predators were released prophylactically and curatively in separate tunnels when plants were artificially infested with spider mites, and at low and moderate spider mite populations when infestations occurred naturally. Although spider mite populations did not establish well the first year, fewer spider mites were recovered with release of N. californicus than with all other treatments. In the second year, spider mites established and the prophylactic release of N. californicus compared favorably with the acaricide-treated plants. In the course of monitoring arthropod populations, we observed a significant reduction in western flower thrips (Frankliniella occidentalis) populations in tunnels treated with N. californicus as compared with non-treated control tunnels. Our field trials validate results obtained from potted-plant experiments and confirm that N. californicus is a superior spider mite predator at high temperatures and low humidities.     

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.     

Low-Density Releases of Neoseiulus Fallacis Provide for Rapid Dispersal and Control of Tetranychus Urticae (Acari: Phytoseiidae, Tetranychidae) on Apple Seedlings

Releases of Neoseiulus fallacis (Garman) at 1500--6000 per ha when prey were at 0.1-0.3 per leaf provided seasonal control of Tetranychus urticae Koch (all stages) at 1-2 per leaf in an apple seedling rootstock nursery. Predaceous mites (all stages) increased to 0.3-0.4 per leaf after releases and predator prey ratios of < or = 1:3-7 provided pest regulation thereafter. Such low-density releases were thought to be effective because multiple dispersal bouts allowed predators to locate widely distributed spider mites (on 2-6% of leaves). A random-diffusion model simulating predator dispersal (incorporating wind speed and direction parameters) adequately explained movement and pest control patterns. An upright, dense, uniform planting of apple seedlings was an effective producer and recipient for dispersing predators and these attributes seemed to explain why biological control was so effective. Low-density releases of N. fallacis for control of T. urticae are predicted to be less effective on other crops with less prominent profiles and soil coverage.     

Within-plant distribution of twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), on ivy geranium: development of a presence-absence sampling plan

The twospotted spider mite, Tetranychus urticae Koch, is an important pest of ivy geranium and other ornamental plants. As a part of our long-term goal to develop an integrated crop management program for ivy geraniums, the focus of this study was to produce a reliable sampling method for T. urticae on this bedding plant. Within-plant mite distribution data from a greenhouse experiment were used to identify the young-fully-opened leaf as the sampling unit. We found that 53% of the mites on a plant are on the young-fully-opened leaves. On average 22, 37, and 41% of the leaves belonged to the young, young-fully-opened, and old leaf categories, respectively. We then developed a presence-absence sampling method for T. urticae in ivy geranium using generic Taylor's coefficients for this pest. We found the optimal binomial sample sizes for estimating populations of T. urticae at densities of between 0 and 3 mites/leaf to be quite large; therefore, we recommend the use of numerical sampling within this range of T. urticae densities. We also suggest that population estimates of T. urticae on ivy geranium be done based on mite density/unit area of greenhouse space, both for conventional greenhouse pest management, and for determining how many phytoseid predators to release when using biological control.     

Effect of tomato leaf hairiness on functional and numerical response of Neoseiulus californicus (Acari: Phytoseiidae)

The objective of this study was to determine whether differences in hairiness of tomato plants affect the functional and numerical response of the predator Neoseiulus californicus McGregor attacking the two-spotted spider mite, Tetranychus urticae Koch. Two tomato hybrids with different density of glandular hairs were used. The functional response was measured by offering eggs and adults of T urticae at densities ranging from 4 to 64 items per tomato leaflet (surface ca. 6.3 cm2); eggs were offered to predator protonymphs and deutonymphs, adult spider mites to adult predators. The number of spider mites eaten as a function of initial density was fitted to the disc equation. Predator densities were regressed against initial prey densities to analyze the numerical response. The number of eggs and adults of T. urticae eaten by N. californicus was extremely low in both hybrids. The nymphal stage of N. californicus and prey density had a significant effect on the number of T urticae eggs eaten by the predator, while hybrid had no effect. The functional response fitted reasonably well to the Holling model. The handling time (Th) and the attack rate (a) were very similar among the two hybrids. The numerical response indicated that the absolute density of predators increased with changes in spider mite densities but the relative predator/prey density decreased in both hybrids. Tomato hairiness prevented N. californicus from exhibiting a strong numerical response and the predator functional response was much lower than observed in other host plants and other phytoseiids. This result shows the need to consider plant attributes as an essential and interactive component of biological control practices.     

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.     

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.     

Spider Mites Avoid Plants with Predators

While searching for food, prey can use cues associated with their predators to select patches with a reduced predation risk. In many cases, odours indicate the presence of both food and predators. Spider mites are known to use odours to locate food and mates, but also to avoid interspecific competitors. We studied the response of the two-spotted spider mite, Tetranychus urticae, to cues associated with the presence of their predators, the phytoseiid Phytoseiulus persimilis. We found that the spider mites strongly avoid plants defended by this predator, but do not avoid plants with another predatory mite, Neoseiulus californicus. Since P. persimilis is commonly used in the greenhouse where our strain of T. urticae was collected and strains of this pest are known to adapt to greenhouse environments, we hypothesize that there has been selection on the pest to recognize its enemy. We further hypothesize that there has been no selection to recognize N. californicus, as it has not been used against two-spotted spider mites in the greenhouse where our spider mites were collected. We discuss the implications of avoidance of predation by spider mites and non-lethal effects of predators for biological control of this pest in greenhouses.     

Predatory role of Neoseiulus fallacis (Acari: Phytoseiidae): spatial and temporal dynamics in Washington red raspberry fields

The seasonal abundance of spider mites and their predator Neoseiulus fallacis (Garman) (Acari: Phytoseiidae) was determined during three consecutive years in Washington State red raspberry fields. Tetranychus urticae Koch (Acari: Tetranychidae), Eotetranychus carpini borealis (Ewing) (Acari: Tetranychidae), and N. fallacis were commonly found in Skagit and Whatcom Counties. E. carpini borealis colonized the fruiting canes earlier in the season than T. urticae. The two phytophages overlapped in midseason, but T. urticae entered diapause earlier than E. carpini borealis and N. fallacis. Densities of N. fallacis increased with increase in spider mite densities. However, the numerical response of the predator was more evident for T. urticae than for E. carpini borealis. Nevertheless, the predator was spatially associated with the two prey species. The spatial and seasonal distribution of N. fallacis in relationship to host plant phenology and prey distribution may influence the effectiveness of this predator as a biological control agent against spider mites in red raspberry. Densities of the predator increased too late to prevent spider mite damage. The predatory role of N. fallacis could be enhanced by introducing or conserving predators that are more tolerant to climatic factors that prevail in and around the cane canopy in the beginning of the season.     

The possibilities for control of two‐spotted spider mite Tetranychus urticae on field‐grown strawberries in the UK by predatory mites

The potential of two species of predatory phytoseiid mites to control the two‐spotted spider mite, Tetranychus urticae, on unprotected field‐grown strawberries in the UK was investigated. Typhlodromus pyri, a native species, was able to control T. urticae in some circumstances, but was less effective under hot, dry conditions. Natural colonization by this species is likely to be too slow in plantations of 1–2 years life. Attempts to introduce this predator to strawberry fields on cut apple shoots were unsuccessful. The exotic predator Phytoseiulus persimilis, which is available from commercial sources, gave very good control of T. urticae when released onto strawberry plants in June. A release earlier in the year was less successful, so to prevent early build‐up of mites and possible plant damage it may sometimes be necessary to reduce spider mite numbers early in the year with an acaricide, before a later release of P. persimilis. The success of this integrated control strategy was demonstrated.     

Within-plant distribution of twospotted spider mites (Acari: Tetranychidae) on impatiens: development of a presence-absence sampling plan

The twospotted spider mite, Tetranychus urticae Koch, is an important pest of impatiens, a floricultural crop of increasing economic importance in the United States. The large amount of foliage on individual impatiens plants, the small size of mites, and their ability to quickly build high populations make a reliable sampling method essential when developing a pest management program. In our study, we were particularly interested in using spider mite counts as a basis for releasing biological control agents. The within-plant distribution of mites was established in greenhouse experiments and these data were used to identify the sampling unit. Leaves were divided into three zones according to location on the plant: inner, intermediate, and other. On average, 40, 33, and 27% of the leaves belonged to the inner, intermediate, and other leaf zones, respectively. However, because 60% of the mites consistently were found on the intermediate leaves, intermediate leaves were chosen as the sampling unit. These results lead to the development of a presence-absence sampling method for T. urticae by using Taylor coefficients generic for this pest. The accuracy of this method was verified against an independent data set. By determining numerical or binomial sample sizes for consistently estimating twospotted spider mite populations, growers will now be able to determine the number of predatory mites that should be released to control twospotted spider mites on impatiens.     

Effects of dispersal, predators (Acari: Phytoseiidae), weather, and ground cover treatments on populations of Tetranychus urticae (Acari: Tetranychidae) in apple orchards

In a 2-yr study of causes of mite outbreaks in apple (Malus spp.) orchards in Nova Scotia, we monitored immigration of Tetranychus urticae Koch from orchard ground cover into trees populated by the generalist phytoseiid predator Typhlodromus pyri Scheuten. In both years, T. urticae-days in the tree canopy increased with number of T. urticae caught in sticky bands on tree trunks. In 2000, T. urticae-days were negatively correlated with T. pyri-days. Lack of correlation in 2001 was attributed to higher rates of immigration, which would mask the effects of predation. Weather also affected mite dynamics. Rainfall in July and August was less in 2001 than in 2000. Heat units were sufficient for six generations of T. urticae in 2001 but only for five in 2000. Consequently, T. urticae-days in the tree canopy and immigration rates were significantly greater in 2001 than in 2000, despite three-fold greater use of miticides. We also tested the effects of herbicides on T. urticae immigration. Application of selective herbicides in laneways reduced coverage of reproductive hosts of T. urticae, but these changes did not reduce immigration. In 2001, application of a miticidal herbicide, glufosinate, in tree rows reduced captures of T. urticae on sticky bands in high immigration orchards but not in low immigration orchards. We conclude that generalist predators and modified herbicide use are insufficient remedies and that effective biological control of T. urticae in the ground cover by a specialist phytoseiid such as Amblyseius fallacis Garman is essential to prevent outbreaks.     

苹果园二斑叶螨种群的空间格局

种群空间格局的研究是昆虫生态学的重要内容,它不仅揭示出种群的空间结构特征,而且还是确定抽样技术和资料代换的基础,二斑叶螨是苹果园的重要害螨,应用4种聚集度指标和Iwao法分别考查了该螨在苹果树内的空间格局及动态规律,结果表明,二斑叶螨在树内不同方向和高度上均以个体群的形式存在,个体群的分布为聚集分布,其中上层和南面树冠的聚集度最高,而下层和内部树冠的聚集度最低,造成这种差异与该螨的生物学特性和环境条件的异质性有关,不论螨体在上层,中层或下层树冠,都明显地表现出前期高聚块,6月中旬以后聚集强度逐渐降低的趋势。     

Sequence analysis of the ribosomal internal transcribed spacers region in spider mites (Prostigmata: Tetranychidae) occurring in citrus orchards in Eastern Spain: use for species discrimination

Tetranychus urticae is a polyphagous mite which is an important pest of citrus worldwide. This mite can be found feeding on many plant species occurring in the citrus agrosystem moving from weeds to trees. Because field samples consist of a mixture of different Tetranychidae species, as a first step necessary to further implement population characterisation of T. urticae, species‐discriminating criteria based on molecular techniques are needed. In this study, the nucleotide variation of the internal transcribed spacers (ITS) 1 and 2 and the intergenic 5.8S fragment of nuclear rDNA of T. urticae, Tetranychus turkestani, Tetranychus evansi, Tetranychus ludeni and Panonychus citri have been determined. Results demonstrate that for these species, the rDNA ITS2 regions are much more conserved than the corresponding rDNA ITS1. The high homogeneity of the ITS2 sequence observed among the specimens of T. urticae obtained from the same ecoregion makes this DNA sequence an excellent tool for species discrimination. ITS sequences differentiate not only species but also specimens from different geographical origin. Furthermore, polymerase chain reaction–restriction fragment length polymorphism analysis of the ITS2 proved adequate for a quick screening of high numbers of field samples.     

Effects of NaCl-stressed citrus plants on life-history parameters of Tetranychus urticae (Acari: Tetranychidae)

Tetranychus urticae is an important pest of citrus, especially lemon and mandarin, under Mediterranean climate. Factors leading to this problem are poorly understood, but saline stress is suspected to contribute to spider mite outbreaks. In this study, the effect of NaCl concentration in nutritive solutions used to water potted young mandarin trees on population growth of T. urticae reared on leaf discs obtained from these plants was investigated. Although the differences observed between treated (5, 10, 30 and 60 mM NaCl) and control groups were in most cases not significant, when all biological parameters calculated were combined to obtain Ro, T and r(m), remarkable differences appeared, and a concentration-dependent effect was detected. Although high salt concentrations negatively affected T. urticae, at the lowest concentration tested the r(m) value was significantly higher than at the water control and this may contribute to the observed field explosions of T. urticae.     

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.     

The role of prey taxis in biological control: a spatial theoretical model

We study a reaction-diffusion-advection model for the dynamics of populations under biological control. A control agent is assumed to be a predator species that has the ability to perceive the heterogeneity of pest distribution. The advection term represents the predator density movement according to a basic prey taxis assumption: acceleration of predators is proportional to the prey density gradient. The prey population reproduces logistically, and the local population interactions follow the Holling Type II trophic function. On the scale of the population, our spatially explicit approach subdivides the predation process into random movement represented by diffusion, directed movement described by prey taxis, local prey encounters, and consumption modeled by the trophic function. Thus, our model allows studying the effects of large-scale predator spatial activity on population dynamics. We show under which conditions spatial patterns are generated by prey taxis and how this affects the predator ability to maintain the pest population below some economic threshold. In particular, intermediate taxis activity can stabilize predator-pest populations at a very low level of pest density, ensuring successful biological control. However, very intensive prey taxis destroys the stability, leading to chaotic dynamics with pronounced outbreaks of pest density.     

Approaches for sampling the twospotted spider mite (Acari: Tetranychidae) on clementines in Spain

Tetranychus urticae Koch (Acari: Tetranychidae) is an important pest of clementine mandarins, Citrus reticulata Blanco, in Spain. As a first step toward the development of an integrated crop management program for clementines, dispersion patterns of T. urticae females were determined for different types of leaves and fruit. The study was carried out between 2001 and 2003 in different commercial clementine orchards in the provinces of Castelló and Tarragona (northeastern Spain). We found that symptomatic leaves (those exhibiting typical chlorotic spots) harbored 57.1% of the total mite counts. Furthermore, these leaves were representative of mite dynamics on other leaf types. Therefore, symptomatic leaves were selected as a sampling unit. Dispersion patterns generated by Taylor's power law demonstrated the occurrence of aggregated patterns of spatial distribution (b > 1.21) on both leaves and fruit. Based on these results, the incidence (proportion of infested samples) and mean density relationship were developed. We found that optimal binomial sample sizes for estimating low populations of T. urticae on leaves (up to 0.2 female per leaf) were very large. Therefore, enumerative sampling would be more reliable within this range of T. urticae densities. However, binomial sampling was the only valid method for estimating mite density on fruit.