Biotic resistance limits the invasiveness of the western flower thrips,Frankliniella occidentalis (Thysanoptera: Thripidae), in Florida

The spread of the western flower thrips, Frankliniella occidentalis (Pergande), has resulted in the world‐wide destabilization of established integrated pest management programs for many crops. It is hypothesized that frequent exposure to insecticides in intensive agriculture selected for resistant populations, which allowed invasive populations in the eastern USA to overcome biotic resistance from the native community of species. Research conducted in Florida to understand the role of biotic factors in limiting the abundance of the western flower thrips is reviewed. Orius spp. (Hemiptera: Anthocoridae) are effective predators that suppress populations of thrips on crop and non‐crop hosts in southern and northern Florida. Orius are more effective predators of the western flower thrips than the native flower thrips, F. tritici (Fitch) and F. bispinosa (Morgan). The native species are competitors of the western flower thrips. Excessive fertilization and the use of broad‐spectrum insecticides in crop fields further enhances populations of the western flower thrips. Interactions with native species clearly limit the abundance of western flower thrips in Florida, but populations are abundant in fertilized crop fields where application of insecticides excludes predators and competitor species.     

Differential predation by the generalist predator Orius insidiosus on congeneric species of thrips that vary in size and behavior

We investigated interactions between the generalist predator Orius insidiosus (Say) (Heteroptera: Anthocoridae) and two species of thrips prey, Frankliniella bispinosa (Morgan) and Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and interspecific differences in morphology and behavior between these prey species that could contribute to differences in predation by O. insidiosus. Frankliniella occidentalis is significantly larger than F. bispinosa. Frankliniella bispinosa has greater mobility compared with F. occidentalis. When O. insidiosus was offered either F. bispinosa or F. occidentalis as prey in single species trials, there were no significant differences in the number of prey captured. However, O. insidiosus had significantly more encounters with F. bispinosa than with F. occidentalis. In arenas with equal numbers of both species, O. insidiosus encountered and captured F. occidentalis more than F. bispinosa. In large arenas with two pepper plants (Capsicum annuum L.), O. insidiosus preyed on more F. occidentalis than on F. bispinosa. These results indicate that O. insidiosus can prey on both thrips species, but that it preferentially captures F. occidentalis. The greater locomotion and movement of F. bispinosa, perhaps combined with its smaller size, allow it to evade predation by O. insidiosus better than F. occidentalis. Consequently, the observed preference of O. insidiosus for F. occidentalis is not exclusively a function of active selection by the predator but also could arise from inherent differences among prey. We propose this differential predation as a mechanism contributing to observed differences in the temporal dynamics of these species in pepper fields.     

Does natural deposition of pine pollen affect the ovipositional behavior of Frankliniella occidentalis and Frankliniella fusca?

Tree pollen, especially Pinus spp. (Pinaceae), is shed in large quantities every spring in North America. Pine pollen deposition onto leaves was found to significantly influence the ovipositional behaviors of certain thrips species (Thysanoptera: Thripidae) in peanut and tomato leaf choice and no‐choice tests. Pine pollen (Pinus elliottii Engelm.) increased the oviposition rate 2.9‐fold for Frankliniella occidentalis (Pergande) (western flower thrips) and 1.6‐fold for Frankliniella fusca (Hinds) (tobacco thrips) in choice tests averaged over both plant species. These results support the idea that pollen has a greater impact on F. occidentalis behavior than on F. fusca behavior. The most dramatic increase was in peanut, where F. occidentalis only oviposited on leaves dusted with pollen, suggesting that the addition of pollen stimulated this flower thrips to lay eggs on a poor host‐plant part. The impact of pollen on the rate of oviposition by thrips is important because it is the early‐instar nymphs that acquire tomato spotted wilt virus (TSWV), which these two thrips species vector. In a laboratory bioassay, the addition of pine pollen to TSWV‐infected peanut foliage increased the percentage of infected F. fusca after one generation.     

Rapid diagnosis of the invasive species,Frankliniella occidentalis (Pergande): a species‐specific COI marker

The western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), is an invasive species and currently occurs in only a few areas in China. An easy, accurate and developmental‐stage independent method to identify F. occidentalis would be a valuable tool to facilitate pest management decision making and, more importantly, to provide an early warning so actions can be taken to prevent its introduction into non‐infested areas. Morphological identification of thrips adults and, to a lesser extent, of second‐stage larvae is the main method currently available to identify F. occidentalis. Molecular identification, however, can be easily carried out by a non‐thrips‐specialist with a little training. In this study, DNA sequence data [within the mitochondrial cytochrome oxidase I gene (COI)] and polymerase chain reaction (PCR) were utilized to develop a molecular diagnostic marker for F. occidentalis. A primer set and PCR cycling parameters were designed for the amplification of a single marker fragment (340 bp) of F. occidentalis COI mtDNA. Specificity tests performed on 28 thrips species, efficacy tests performed on five immature developmental stages as well as on male and female adults and tests on primer sensitivity all demonstrated the diagnostic utility of this marker. Furthermore, the primer set was tested on seventeen F. occidentalis populations from different countries and invaded areas in China and proved to be applicable for all geographic populations. It was used successfully to clarify the distribution of F. occidentalis in the Beijing metro area. These results suggested that this diagnostic PCR assay provides a quick, simple and reliable molecular technique for the identification of F. occidentalis.     

Behavioral Response of Western Flower Thrips to Visual and Olfactory Cues

Behavioral responses of Frankliniella occidentalis Pergande (Thysanoptera: Thripidae) to visual and olfactory cues were assessed in a cylindrical black box and a Y-tube olfactometer. Frankliniella occidentalis preferred circular shapes to other five geometrical patterns (rectangle, triangle, inverted triangle, diamond, and modified circle imitating flower petals) in multiple choice tests. In pair wise choice tests, the thrips preferred the yellow artificial flower shape to the geometrical patterns tested. Frankliniella occidentalis stayed on the artificial flower about four times longer than on the geometrical patterns. Higher numbers of thrips responded to a combination of p-anisaldehyde and artificial flower compared to the arm with only the olfactory or the visual cue. These results suggest that shape is an important cue for F. occidentalis.     

Continuous exposure to the deterrents cis‐jasmone and methyl jasmonate does not alter the behavioural responses of Frankliniella occidentalis

Behavioural responses of Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), a generalist, cell sap‐feeding insect species with piercing‐sucking mouthparts, after continuous exposure to two deterrent secondary plant compounds are investigated. We compared in choice assays on bean leaf discs, the settling, feeding, and oviposition preferences of F. occidentalis females that had no experience with the two fatty acid derivatives methyl jasmonate and cis‐jasmone before testing (naïve thrips) vs. females that had been exposed to the deterrent compounds before testing (experienced thrips). The thrips were exposed to the deterrents at low or high concentrations for varied time periods and subsequently tested on bean leaf discs treated with the respective deterrent at either a low or a high concentration. Frankliniella occidentalis females avoided settling on the deterrent‐treated bean leaf discs for an observation period of 6 h, independent of their previous experience. Our results demonstrate that feeding and oviposition deterrence of the jasmonates to the thrips were not altered by continuous exposure of the thrips to the jasmonates. Habituation was not induced, neither by exposure to the low concentration of the deterrents nor by exposure to the high concentration. These results indicate that the risk of habituation to two volatile deterrent compounds after repeated exposure is not evident in F. occidentalis. This makes the two compounds potential candidates to be integrated in pest management strategies.     

Prey suitability of western flower thrips,Frankliniella occidentalis,and onion thrips,Thrips tabaci,for the predatory mite Amblyseius swirskii

Western flower thrips, Frankliniella occidentalis, and onion thrips, Thrips tabaci, are both important polyphagous pests of vegetables and ornamentals in greenhouses. Difficulties in biological control of these pests have prompted a search for new natural enemies. Most recently, the predatory mite Amblyseius swirskii has been commercialised as biological control agent of whiteflies and thrips. However, little is known about the suitability of thrips as prey for A. swirskii. We therefore assessed prey acceptance and life history of A. swirskii when feeding on F. occidentalis and T. tabaci at 25±1°C. Amblyseius swirskii juveniles preyed upon first larval instars of both F. occidentalis and T. tabaci but suffered from high mortality (67 and 78%). Developmental time (egg to adult) of A. swirskii was 7.8 days with either prey species. Adult A. swirskii females readily accepted first larval instars of both thrips species, which were attacked in <20 min on a leaf and <10 min in an artificial cage. Oviposition rates (0.92 and 0.99 eggs/female/day) and offspring sex ratios (63 and 70% females) were similar with F. occidentalis and T. tabaci as prey. Less than one-third of juveniles reaching adulthood and oviposition rates below one egg/female per day resulted in relatively low intrinsic rates of increase (r _m) (0.056 and 0.024 per day with F. occidentalis and T. tabaci, respectively). Altogether, our study suggests that the recently reported superiority of A. swirskii to the widely used Neoseiulus cucumeris in suppression of thrips is due to other traits than its population growth capacity with thrips as prey.     

Population abundance and movement of Frankliniella species and Orius insidiosus in field pepper

1 A recent study revealed the capacity of the Orius insidiosus to suppress populations of Frankliniella spp. in field pepper during the spring when thrips are rapidly colonizing and reproducing. In this study, population abundance in pepper during spring, summer, and autumn was determined to understand better predator/prey dynamics under local conditions. Local movement between pepper flowers also was quantified to examine how population attributes of the predator allow suppression of rapidly moving populations of prey. 2 Randomized complete block experiments established in the autumn of 1998 and the spring of 1999 included treatments of biological and synthetic insecticides, which altered the population densities of predator and prey. Numbers of O. insidiosus in relation to prey were sufficient in 1998 to prevent build‐up of thrips populations. In 1999, populations of thrips were unable to recover from near extinction owing to persistence of the predator. The predator rapidly recolonized plots treated with insecticide. 3 Greenhouse plants of the same age as field plants were used to monitor movement by predators and prey. Movement by F. occidentalis was limited, whereas F. tritici and F. bispinosa moved rapidly to the greenhouse plants. The males of each thrips species moved more rapidly than the females. There was evidence that rapid movement assisted F. tritici and F. bispinosa in avoiding predation, but O. insidiosus also moved very rapidly to the greenhouse plants. This attribute explains the predator's ability to suppress thrips rapidly even when populations are rapidly colonizing and reproducing in the flowers.     

Relationships between Tospovirus Incidence and Thrips Populations on Tomato in Mendoza, Argentina

The objectives of this work were to estimate the capability of local populations of thrips as vectors of groundnut ringspot virus (GRSV) and tomato spotted wilt virus (TSWV), and to determine the species composition of vectors in tomato crops. Transmission assays were performed. Incidence of tospoviruses was estimated in commercial crops. Random samples of flowers were taken from tomato for identification of thrips. Of the five species of thrips tested, Frankliniella gemina (first record), F. occidentalis and F. schultzei transmitted GRSV and TSWV. F. schultzei was a significantly more efficient vector of GRSV than F. occidentalis under controlled assay conditions. The thrips were identified on flowers from six surveyed tomato crops. F. occidentalis was the most frequently identified species (43.0%), followed by F. schultzei (35.6%) and Thrips tabaci (10.1%). The incidence of tospoviruses was low (1.1–2.8%) in crops planted during August–September and greater (9.5–61.1%) in crops planted in December. GRSV was prevalent (85%) over TSWV (11%).     

Variation in thrips species composition in field crops and implications for tomato spotted wilt epidemiology in North Carolina

Thrips were surveyed in tomato spotted wilt-susceptible crops in five areas across North Carolina. Tomato, pepper, and tobacco plants in commercial fields were sampled and 30 species of thrips were collected over a 3-year period. The most common species overall was Frankliniella tritici (Fitch). The most common thrips species that are known to vector Tomato Spotted Wilt Virus (TSWV) were F. fusca (Hinds), and F. occidentalis (Pergande). Relatively low numbers of Thrips tabaci Lindeman, another reported vector, were collected. The spatial and temporal occurrence of vectors varied with sampling method, crop species, region of North Carolina, and localized areas within each region. In a laboratory experiment, no difference was detected between the ability of F. fusca and F. occidentalis to acquire and transmit a local isolate of TSWV. Based on vector efficiency and occurrence, F. fusca is considered the most important vector of TSWV in tobacco, whereas both F. fusca and F. occidentalis are important vectors of TSWV in tomato and pepper.     

西花蓟马高毒力虫生真菌筛选及其生防应用潜力的研究进展

西花蓟马是一种外来入侵的世界性害虫,对农林业危害巨大。查阅国内外相关文献,综述了当前防治西花蓟马的虫生真菌的种类、高毒力菌株的筛选及防治现状。现已知西花蓟马的寄生病原真菌有5种,包括蜡蚧轮枝菌(半知菌:丝孢目)、球孢白僵菌(半知菌:丝孢目)、金龟子绿僵菌(半知菌:丝孢目)、玫烟色棒束孢(半知菌:束梗孢目)和小孢新接霉。其中,球孢白僵菌、金龟子绿僵菌在西花蓟马的生物防治中应用最广,具有良好的开发应用潜力,部分防效好的虫生真菌已申请专利及实现工厂化生产。     

Life history characteristics of Frankliniella occidentalis on cucumber leaves with and without supplemental food

The development time, fecundity, longevity, and resultant intrinsic growth rate of the western flower thrips Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidae] encaged on a cucumber leaf were compared among seven types of food supplied: six pollen species and a mixture of milk powder and yeast. The rationale was to find a food source that offers the least benefit for thrips and could therefore be considered as a food source for the preventative introduction of thrips predators. With the exception of the mixture of milk powder and yeast, all the food sources tested offered a nutritional benefit for the thrips. The addition of pollen increased fecundity and reduced development time, mainly during the larval stage. Betula pubescens and Corylus avellana pollen also increased adult longevity. The nutritional benefit of Pinus sylvestris pollen was greater than that of the other five pollen species, as manifested by its significantly greater positive effect on fecundity. The other pollen species could not be ranked in terms of nutritional benefit to F. occidentalis. The negative effect of the milk powder plus yeast mix on the life‐table parameters of F. occidentalis probably only occurs in an encaged situation where thrips cannot escape from the unfavorable environment. The crude estimate of the intrinsic growth rate of F. occidentalis increased from 0.163 on the plain cucumber leaf to 0.240 when P. sylvestris pollen was added to the leaf. The differences in intrinsic growth rate mainly reflect the differences in fecundity among the food sources. Thus, the peak oviposition rate may be used as a measure of the nutritional benefit F. occidentalis can obtain by feeding on supplemental food sources. The positive effect of a supplemental food source on thrips does not necessarily mean it is unsuitable for the preventative introduction of thrips predators, because the supplemental food can also affect the population dynamics of the predator and the predator–prey interaction and, hence, the outcome of biological control.     

Use of spinosad and predatory mites for the management of Frankliniella occidentalis in low tunnel‐grown strawberry

Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) is a major pest of strawberry. The efficacy of three species of predatory mites, Typhlodromips montdorensis (Schicha), Neoseiulus cucumeris (Oudemans) (both: Acari: Phytoseiidae), and Hypoaspis miles (Berlese) (Acari: Laelapidae), and their compatibility with spinosad for the control of this thrips was evaluated in commercial strawberry in spring. Low tunnel‐grown strawberry was treated with ‘water then mites’, ‘spinosad then mites’, or ‘mites then spinosad’. Predatory mites were released as single‐, two‐ and three‐species combinations. Overall, spinosad‐treated plants had fewer thrips than water‐treated plants (control). In all treatment regimes, each species of predatory mite reduced the number of thrips relative to those plants that received no mites. Predatory mites were most effective in reducing thrips when released after spinosad was applied. Any multiple‐species combination of predatory mites reduced thrips numbers more than single‐species releases. The two‐species combination of T. montdorensis (foliage inhabiting) and H. miles (soil dwelling) was the most effective in suppressing thrips. The next most effective combination was a three‐species release. Of multiple‐species combinations, the two‐species combination of T. montdorensis and N. cucumeris was the least effective in suppressing thrips numbers. The spinosad and mites only temporarily reduced the numbers of F. occidentalis. This suggests that further application of predatory mites, spinosad, or both is required to maintain F. occidentalis populations below an economically damaging level.     

Influence of plant species and plant growth stage on Frankliniella occidentalis pupation behaviour in greenhouse ornamentals

The influence of plant stage and plant species on the pupation behaviour of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), was investigated in flowering and non‐flowering potted chrysanthemums and mini roses. On non‐flowering chrysanthemums and roses, 92–93% of the F. occidentalis pupated in the soil. On flowering roses, 87% pupated in the soil, and on flowering chrysanthemums only 60% chose the soil as a pupation site and 40% stayed on the plant. This means that, in the presence of flowers, especially complex inflorescences like chrysanthemums, a large proportion of F. occidentalis chooses stay on the plant to pupate.     

Weed species in tomato production and their role as alternate hosts of Tomato spotted wilt virus and its vector Frankliniella occidentalis

Tomato spotted wilt virus (TSWV) is an important plant virus that infects a wide range of hosts including weeds making its management difficult. A survey was undertaken to establish the occurrence of weed species in tomato production systems in Kenya and their role as hosts of TSWV and its vectors. Selected weed species were further evaluated for their reaction to TSWV, transmission efficiency by Frankliniella occidentalis and ability to support thrips reproduction. Of the 43 weed species identified in the field, 29 species had been reported as hosts of TSWV, two were non‐hosts and 11 had no record of their status. Among the more common species, Amaranthus hybridus, Solanum nigrum, Tagetes minuta and Datura stramonium were susceptible to the virus and supported high levels of thrips reproduction. The TSWV could not be transmitted to Galinsoga parviflora and Sonchus oleraceus by F. occidentalis despite them being highly susceptible in mechanical transmission tests. There was a significant correlation between feeding damage and number of larvae of F. occidentalis on different weeds. Occurrence of weeds that support thrips reproduction and are good hosts of TSWV is a clear indicator of their role in epidemiology and the importance of their management for disease control.     

Effect of intercrops on thrips species composition and population abundance on French beans in Kenya

The study aimed at determining thrips species composition and thrips population density on French bean planted as a sole crop and as an intercrop with either sunflower, Irish potato, or baby corn, in various combinations. Field experiments were conducted in two seasons to examine: (1) thrips population development and thrips species composition over time, (2) effect of intercrops on thrips population density and natural enemies, and (3) effect of intercrops on French bean yield. The experiments were conducted at the Kenya Agricultural Research Institute, Embu, Kenya in a randomized complete block design with four replicates. The thrips population on French beans increased with time. It showed a peak at the flowering stage then started declining when the crops were nearing senescence. French beans hosted four thrips species, Megalurothrips sjostedti (Trybom), Frankliniella schultzei (Trybom), Frankliniella occidentalis (Pergande), and Hydatothrips aldolfifriderici (Karny) (all Thysanoptera: Thripidae) in order of decreasing abundance. The main thrips species on Irish potato and sunflower was F. schultzei. Baby corn hosted only Frankliniella williamsi (Hood) and Thrips pusillus (Bagnall). A monocrop of French bean hosted more thrips than a French bean intercrop mix. Thrips natural enemies such as Orius spp. and Ceranisus spp. were recorded in all crop plants but in especially high numbers on French bean and baby corn, respectively. Plots with French bean alone had about 1.4 times higher yields compared to intercropped plots of French bean with sunflower and French bean with baby corn. However, the percentage of pods that could get rejected on the market due to thrips damage was highest on plots with French bean alone (68 and 63%) and lowest on plots with French bean and baby corn (35 and 37%) in the first and second seasons, respectively. This study showed that a complex of thrips is found in the field and its composition varies with crop stage and species. Intercropping French bean with other crops compromises on French bean yield but reduces damage to the French bean pods, thereby enhancing marketable yield.     

Life-history parameters of western flower thrips on susceptible and resistant cucumber genotypes

The life-history parameters reproduction rate, developmental time and age specific survival of the western flower thrips,Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidae], were determined on susceptible and resistant cucumber (Cucumis sativus L.) genotypes. Both newly emerged andF. occidentalis females of mixed ages showed a substantial reduction (36 to 50%) of the reproduction rate on all resistant genotypes, in particular after the second day. On the resistant genotypes 9127 and 9140,F. occidentalis had a prolonged developmental period. This was primarily due to a prolongation of the second larval stage. On all resistant genotypes,F. occidentalis suffered from high (82 to 97%) preadult mortality, predominantly at the second larval stage. It is conclude that the resistant genotypes do not cause an immediate intoxication of adult nor preadult thrips stages.     

Distributions of western flower thrips (Thysanoptera: Thripidae) and its predatory bug Orius niger (Hemiptera: Anthocoridae) assessed by coloured sticky traps and plant samplings in cotton

The capturing efficiency of four coloured (yellow, green, white and blue) sticky traps, placed at the top, middle and bottom strata of cotton plants, was tested for the western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and its predatory bug, Orius niger (Wolff) (Hemiptera: Anthocoridae), as well as spatial distributions of both insects on the plant in years 2006 and 2007. The white coloured trap was the most attractive to F. occidentalis and O. niger in the 2-year study. The blue coloured trap was the least attractive for Orius. The mean numbers of F. occidentalis and O. niger on the top plant parts (flowers and leaves) and in all coloured traps positioned on the upper parts of the plants were greater than those found in the lower two strata. Taylor's power law analysis showed that F. occidentalis and O. niger adults aggregated in the flowers or on the leaves. This study suggests that top flowers could be preferentially sampled to determine population densities of Frankliniella flower thrips and Orius species in cotton, and thus, sticky traps should be placed on the top level of plants. F. occidentalis: O. niger ratios in the flowers varied from 4 to 60 thrips per Orius adult in the three plant strata. This result may indicate that F. occidentalis experiences more predation from Orius.     

Predation capacity of two predatory laelapid mites on soil-dwelling thrips stages

The life cycle of the Western Flower Thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), one of the most important glasshouse pests, includes a soil passage composed of three instars that deserve more attention in terms of biocontrol strategies. It has been repeatedly reported that two polyphagous predatory mites, Stratiolaelaps miles (Berlese) and Hypoaspis (Geolaelaps) aculeifer (Canestrini) (Acari: Laelapidae), also prey on these thrips stages, in addition to several other soil inhabiting prey species. However, the potential thrips consumption rates have never been quantified for these predatory mites. Therefore, an arena experiment was carried out to investigate the potential predation rates of the two mites on second instar larvae, prepupae, and pupae of F. occidentalis. In addition, the fecundity on the thrips diet was assessed and compared to oviposition rate on a nematode prey. All thrips instars were accepted as prey by each mite species. Females of H. aculeifer preyed on 3.5 (± 0.5) thrips instars and laid 2.5 (± 0.87) eggs per day, whereas females of S. miles preyed on 1.64 (± 0.3) thrips and laid 0.8 (± 0.53) eggs. Males of both species killed 0.6 (± 0.3) thrips per day. The fitness of the two predatory mites on F. occidentalis as prey and their suitability as biocontrol agents are elucidated. Reasons for reduced thrips control in the soil environment, in contrast to the results obtained in arena assays are discussed.     

Is Xysticus kochi (Araneae: Thomisidae) an efficient indigenous biocontrol agent of Frankliniella occidentalis (Thysanoptera: Thripidae)?

According to the present practice of Hungarian greenhouse sweet pepper production, only exotic agents are used for biological control purposes against thrips pests. The suitability for biological control of the second instars of an indigenous species, the common crab spider, Xysticus kochi Thorell (Araneae: Thomisidae) was tested on thrips species, Frankliniella occidentalis (Pergande) and Frankliniella intonsa (Trybom) (Thysanoptera: Thripidae) using a cage set-up on greenhouse sweet pepper plants. Effects of introducing second instars of X. kochi on thrips infested plants were judged by assessing the degree of aesthetical damage, the commercial value of the fruits (degree of economic loss) and the quality composition of the harvested peppers. The estimated damaged surface unit was significantly lower in control plants than in plants infested by F. occidentalis. A significant effect of the spider introduction was observed in all of the pepper quality indicators applied. We suspect that direct Xysticus predation or any other predator-induced effect resulted in reduced levels of damage on harvested peppers. However, further investigation is needed to detect the origin (e.g., predation and non-predation effects) of the suppression of thrips damage. Our results suggest that X. kochi could be an effective component of greenhouse antagonistic fauna and the application of mulch may encourage the effectiveness of spiderlings. An erratum to this article can be found at