Impact of production system on development of insecticide resistance in Frankliniella occidentalis (Thysanoptera: Thripidae)

The western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), has become one of the most difficult insects to control in the intensive agriculture of southeastern Spain. However, resistance problems are quite different in two neighboring areas, Murcia and Almeria, with distinct production systems. Thirty-six field populations of western flower thrips from sweet pepper crops were collected in two different dates in Murcia and Almeria in 2005 and 2006. Western flower thrips populations collected were exposed to a diagnostic concentration of spinosad, methiocarb, acrinathrin, and formetanate. The results allowed the recognition of higher levels of resistance in Almeria compared with Murcia throughout the growing season. The mortality at the diagnostic concentration for spinosad (120 ppm) in western flower thrips populations ranged from 34 to 81% in Almeria, and from 73 to 100% in Murcia. The mortalities at the diagnostic concentration to acrinathrin (800 ppm) and formetanate (8000 ppm) were 17-31% in Almeria and 77-100% in Murcia, and 14-41% in Almeria and 48-99% in Murcia, respectively, indicating large geographic variations. Toxicity of methiocarb was higher for western flower thrips populations from both areas. However, mortality at the diagnostic concentration of methiocarb (2000 ppm) varied from 56 to 90% in Almeria, and it was from 94 to 100% in Murcia. The impact of production systems and agricultural practices of each area on the development and stability of insecticide resistance is discussed.     

Mechanisms associated with methiocarb resistance in Frankliniella occidentalis (Thysanoptera: Thripidae)

Biochemical mechanisms associated with methiocarb resistance were examined in laboratory-selected and field populations of the western flower thrips, Frankliniella occidentalis (Pergande). Seven populations were examined and they differed in their susceptibility to methiocarb by 30 times. Including the synergists piperonyl butoxide, a cytochrome P-450 monooxygenase inhibitor, or S,S,S-tributylphosphorotrithioate, an esterase inhibitor, in the methiocarb bioassays partially suppressed resistance in the most resistant populations. In vitro assays of general esterase, glutathione S-transferase, and acetylcholinesterase activities showed increased activity in some of the resistant populations and increased activity of the enzymes after methiocarb selection on one of the populations. Assays of acetylcholinesterase sensitivity to inhibition by methiocarb, dichlorvos, and eserine suggested insensitive acetylcholinesterase in two of the resistant populations. These results indicate that methiocarb resistance in F. occidentalis was polyfactorial and involved detoxification and altered target site. None of the biochemical assays showed interpopulation enzymatic differences strongly correlated with the level of methiocarb resistance. The possibilities for developing rapid biochemical diagnostic assays to detect methiocarb resistance in F. occidentalis are discussed.     

Life-stage variation in insecticide resistance of the western flower thrips (Thysanoptera: Thripidae)

The life-stage variations in insecticide resistance of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), to selective insecticides (acrinathrin, formetanate, and methiocarb) were studied using resistant laboratory strains. In each strain, the second-instar larva was less susceptible to the insecticides tested than the adults. The lower the resistance level of the adults, the higher the difference between larva and adult susceptibility: 32-fold to methiocarb, 15.4-fold to formetanate, and 180-fold to acrinathrin in the reference strain. In laboratory-selected resistant strains, these differences were much lower: 5.8-fold to methiocarb, 4.8-fold to formetanate, and 2.0-fold to acrinathrin. In selected strains, higher resistance levels for each insecticide were found, both for larvae and adults, compared with the reference strain. These results show that after insecticide resistance selection in adults, the resistance is carried over to the larvae, but at lower levels.     

Management of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), in hydroponic strawberry crops: using yellow sticky traps to determine action thresholds

Abstract  The relationship between western flower thrips (WFT) populations in flowers and catches on yellow sticky traps was investigated in a hydroponic strawberry crop in the Sydney area in 1999–2000. The thrips population was initially dominated by males, but by mid-summer it was composed primarily of females. At a point when approximately 60–65% of WFT on traps were female (approximately 20–30 females per trap), the overall density of thrips on traps and in the crop increased rapidly, resulting in severe damage to immature fruit. The sex ratio of 0.60–0.65 females corresponded very closely to a mean of five adult WFT per flower at the start of the trapping week, which was an infestation level previously calculated as the action threshold to prevent damage to young green fruit.     

Control of Western flower thrips (Frankliniella occidentalis Pergande) pupae in compost

Western flower thrips (Frankliniella occidentalis) develops from an active larval stage through to a non-feeding, almost immobile, pre-pupal and pupal stage. This generally occurs in the compost or soil below the plant on which the larvae fed. Control of thrips at this stage in their development offers a chance of utilising pathogens or pesticides not suitable for use in an integrated control programme aimed at adult and larval stages. Trials were done with F. occidentalis pupae and pre-pupae in a soil/peat based compost using 11 pesticides, three fungal pathogens and four species of insect parasitic nematodes. The pesticides malathion and chlorpyrifos-methyl gave the most promising result with 97.5% and 96.5% control, respectively. The fungal pathogen Metarhizium anisopliae proved better when applied as a pre-pupation rather than as a post-pupation treatment (74.5% : 26.6% control). The insect parasitic nematode Steinernema carpocapsae gave 76.6% control when applied at 25 times 10⁴ nematodes litre^-1 of compost. The results are discussed in relation to control of thrips in glasshouses.     

Population dynamics of thrips prey and their mite predators in a refuge

Prey refuges are expected to affect population dynamics, but direct experimental tests of this hypothesis are scarce. Larvae of western flower thrips Frankliniella occidentalis use the web produced by spider mites as a refuge from predation by the predatory mite Neoseiulus cucumeris. Thrips incur a cost of using the refuge through reduced food quality within the web due to spider mite herbivory, resulting in a reduction of thrips developmental rate. These individual costs and benefits of refuge use were incorporated in a stage-structured predator-prey model developed for this system. The model predicted higher thrips numbers in presence than in absence of the refuge during the initial phase. A greenhouse experiment was carried out to test this prediction: the dynamics of thrips and their predators was followed on plants damaged by spider mites, either with or without web. Thrips densities in presence of predators were higher on plants with web than on unwebbed plants after 3 weeks. Experimental data fitted model predictions, indicating that individual-level measurements of refuge costs and benefits can be extrapolated to the level of interacting populations. Model-derived calculations of thrips population growth rate enable the estimation of the minimum predator density at which thrips benefit from using the web as a refuge. The model also predicted a minor effect of the refuge on the prey density at equilibrium, indicating that the effect of refuges on population dynamics hinges on the temporal scale considered.     

The use of triflumuron on sugar-baited targets for autosterilization of the housefly, Musca domestica

A target system, capable of delivering an effective transovarial dose of the chitin synthesis inhibitor triflumuron to the house flyMusca domestica Linnaeus (Diptera: Muscidae), is described. A single 15×5 cm polyester target, dipped in triflumuron 10% suspension concentrate (s.c.) and 50% W/V sucrose solution and suspended in 30×30×30 cm cages of approximately 400 adultM. domestica for 24 h, reduced mean egg hatch to 47%. Egg hatch was reduced to 35% by allowing flies access to similar targets for 7 d. Exposure to triflumuron also interrupted the developmental cycle at the larval stage. After adultM. domestica had been exposed for 24 h to a target baited with 10% triflumuron s.c. and a 50% W/V sucrose solution, only 6% of eclosed larvae pupariated; longer exposure times prevented any pupariation. However, beyond pupariation, there appeared to be no further effect of triflumuron. The age of the flies at first exposure also affected the extent of the ovicidal and larvicidal effects induced, younger adults being more susceptible. However, effects were shown to decrease with time after exposure to triflumuron. A target dosed with 3% triflumuron s.c. resulted in a greater ovicidal effect than a target dosed with 10% triflumuron, suggesting that the higher doses were detected and elicited some negative response from the flies. Effects on females via the treatment of males were demonstrated. The results suggest that utilization of sugar-baited targets dosed with a low concentration of triflumuron may be a useful step in the development of autosterilizing systems for house fly control.     

Predator-prey relationships on Apiaceae at an organic farm

Orius insidiosus (Say) and O. pumilio (Champion) were confirmed to be sympatric in north central Florida as the major predators of the Florida flower thrips, Frankliniella bispinosa (Morgan), on flowers of Queen Anne's lace, Daucus carota L. and false Queen Anne's lace, Ammi majus L. F. bispinosa was the predominant thrips observed on both flowers but colonized D. carota to a greater extent and earlier in the season than A. majus. Despite differences in the abundance of F. bispinosa on the two plants, neither Orius species showed host plant affinities. Population profiles for the thrips and Orius spp. followed a density dependent response of prey to predator with a large initial prey population followed by a rapid decline as the predator populations increased. The temporal increases in Orius spp. populations during the flowering season suggest that they were based on reproductive activity. As observed in a previous study, O. insidiosus had a larger population than O. pumilio and also had a predominantly male population on the flowers. By examining carcasses of the prey, there appeared to be no sexual preference of the thrips as prey by the Orius spp. as the prey pattern followed the demographics of the thrips sex ratio. Few immatures of either thrips or Orius spp. were observed on D. carota or A. majus, which suggests that oviposition and nymphal development occurred elsewhere. Based on these findings, D. carota and A. majus could serve as a banker plant system for Orius spp.     

Jasmonate-dependent plant defenses mediate soybean thrips and soybean aphid performance on soybean

Insect herbivores from different feeding guilds induce different signaling pathways in plants. In this study, we examined the effects of salicylic acid (SA)- and jasmonic acid (JA)-mediated defenses on performance of insect herbivores from two different feeding guilds: cell-content feeders, soybean thrips and phloem feeders, soybean aphids. We used a combination of RT-qPCR analysis and elicitor-induced plant resistance to determine induction of SA and JA signaling pathways and the impact on herbivore performance. In the early interaction between the host plant and the two herbivores, SA and JA signaling seems to occur simultaneously. But overall, soybean thrips induced JA-related marker genes, whereas soybean aphids increased SA and ABA-related marker genes over a 24-h period. Populations of both soybean thrips and soybean aphids were reduced (47 and 25 %, respectively) in methyl jasmonate (MeJA)-pretreated soybean plants. SA treatment has no effect on either herbivore performance. A combination pretreatment of SA and MeJA did not impact soybean thrips population but reduced soybean aphid numbers which was comparable with MeJA treatment. Our data suggest that SA–JA antagonism could be responsible for the effect of hormone pretreatment on thrips performance, but not on aphid performance. By linking plant defense gene expression and elicitor-induced resistance, we were able to pinpoint the role for JA signaling pathway in resistance to two herbivores from different feeding guilds.     

Spatiotemporal population dynamics of the banana rind thrips,Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae)

The understanding of how environmental factors and agricultural practices affect population dynamics of insect pests is necessary for pest management. Here, we provide insight into the ecology of the banana rind thrips Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae) by collecting and analysing a spatiotemporal database of population estimates in Martinique (West French Indies). We assessed the influence of climatic variables (which were rainfall and temperature) and biotic variables (which were banana and three weed species) on the adult thrips abundance for different components of the banana plant (sucker, mother plant and bunch) and evaluated the effect of thrips abundance and standard bunch covers on damages. The abundance of thrips on the sucker, the mother plant, and the bunch was significantly related to the abundance on neighbouring banana plants, and spatial autocorrelation indicated that E. brevisetis dispersed for only short distances. The number of thrips on the mother plant and on the bunch was positively related to the number of thrips on the sucker, suggesting that the thrips may disperse from the sucker to the mother plant and then to the bunch. The abundance of thrips on the sucker increased with sucker height and was positively correlated with the mean daily rainfall during the 17 days before sampling; the length of that period might correspond with the time required for an individual to complete its life cycle. Covered bunches had 98% fewer thrips than non‐covered bunches, and the damage caused by thrips was linearly related to the number of thrips present between the 2nd and 4th week after flowering. Finally, we found that the presence of Alocasia cucullata, Dieffenbachia seguine and Peperomia pellucida is significantly related with a decrease in thrips abundance on banana plants, suggesting the use of these weeds as potential trap plants.     

Sequential sampling of Aphis gossypii Glover (Hemiptera: Aphididae) and Frankliniella schultzei Trybom (Thysanoptera: Thripidae) on cotton crop

The goal of this research was to use the sequential probability ratio test to establish a sequential sampling plan for Aphis gossypii Glover and Frankliniella schultzei Trybom infesting cotton. Field work was conducted at the agricultural experimental station of the Universidade Federal da Grande Dourados during the 2003/2004 and 2004/2005 agricultural years. Aphid colonies and individual thrips in the sampling area were counted and their numbers were recorded. The spatial distribution pattern of A. gossypii and F. schultzei in the cotton culture was aggregated. Sequential sampling plans were developed for aphids and thrips with type I and type II errors set at 0.1, common Kc = 0.6081 (aphids) and = 0. 9449 (thrips), and safety and management levels of 20% (aphids) and 40% (thrips) of infested plants. The sampling plans resulted in two decision boundaries for each species, as follows: the upper boundary, indicating when management (population control) is recommended: S1 = 4.6546 + 0.2849n (aphids), and S1 = 3.6514 + 0.1435n (thrips); and the lower boundary, indicating when population control is not necessary: S0 = -4.6546 + 0.2849n (aphids) and S0 = - 3.6514 + 0.1435n (thrips). The highest probability of error when making a decision was 3% for aphids and 2% for thrips, respectively. The maximum number of samples required to reach a decision was 63 for aphids and 95 for thrips.     

Evaluation of the triflumuron and the mixture of Bacillus thuringiensis plus Bacillus sphaericus for control of the immature stages of Aedes aegypti and Culex quinquefasciatus (Diptera: Culicidae) in catch basins

INTRODUCTION: In Cali, Colombia, catch basins (streetside storm drains) are one of the main larval habitats of Aedes aegypti and Culex quinquefasciatus. Since 1999, these mosquitoes have been controlled by the Secretaría de Salud Municipal (Secretary of Municipal Public Health) using the larvicide triflumuron. Because of high densities of these mosquitoes that remain in the city, treatment failure was suspected -possibly insecticide resistance of the target species. OBJECTIVES: The efficacy of triflumuron and VectoMax (biorational mixture of Bacillus thuringiensis var. israelensis plus Bacillus sphaericus) were evaluated in the control of A. aegypti and C. quinquefasciatus in catch basins. The residual effect of a single application of the biorational formulation was determined in catch basins during periods of high and low rainfall. MATERIALS AND METHODS: The efficacy of the products was measured in 60 catch basins located in a residential neighborhood of Cali for a period of 90 days. The mean number of immature instars (A. aegypti and C. quinquefasciatus larvae and pupae of both species) was determined biweekly from 40 catch basins with insecticide intervention (20 treated with triflumuron, 20 with VectoMax) and 20 untreated (control group). The residual effect of the biorational larvicide was evaluated biweekly in 10 catch basins during each of the 2 climatic periods. Results. The catch basins treated with VectoMax presented a significantly lower mean number of immature instars of both species compared with the control ( p<0.01). In contrast, the triflumuron treatment significantly reduced only immature instars of A. aegypti compared with the control ( p<0.001). The residual effect of VectoMax was higher during low rainfall compared to the control ( p<0.001). Conclusion. The biorational formulation was the more effective treatment for the control of both species during the period of evaluation (15 days).     

Effects on population persistence: the interaction between environmental noise colour,intraspecific competition and space

It is accepted that accurate estimation of risk of population extinction, or persistence time, requires prediction of the effect of fluctuations in the environment on population dynamics. Generally, the greater the magnitude, or variance, of environmental stochasticity, the greater the risk of population extinction. Another characteristic of environmental stochasticity, its colour, has been found to affect population persistence. This is important because real environmental variables, such as temperature, are reddened or positively temporally autocorrelated. However, recent work has disagreed about the effect of reddening environmental stochasticity. Ripa and Lundberg (1996) found increasing temporal autocorrelation (reddening) decreased the risk of extinction, whereas a simple and powerful intuitive argument (Lawton 1988) predicts increased risk of extinction with reddening. This study resolves the apparent contradiction, in two ways, first, by altering the dynamic behaviour of the population models. Overcompensatory dynamics result in persistence times increasing with increased temporal autocorrelation; undercompensatory dynamics result in persistence times decreasing with increased temporal autocorrelation. Secondly, in a spatially subdivided population, with a reasonable degree of spatial heterogeneity in patch quality, increasing temporal autocorrelation in the environment results in decreasing persistence time for both types of competition. Thus, the inclusion of coloured noise into ecological models can have subtle interactions with population dynamics.     

Grass thrips (Anaphothrips obscurus) (Thysanoptera: Thripidae) population dynamics and sampling method comparison in timothy

Sampling studies were conducted on grass thrips, Anaphothrips obscurus (Müller) (Thysanoptera: Thripidae), in timothy, Phleum pratense L. These studies were used to compare the occurrence of brachypterous and macropterous thrips across sampling methods, seasons, and time of day. Information about the population dynamics of this thrips was also revealed. Three absolute and two relative methods were tested at three different dates within a season and three different daily times during four harvest periods. Thrips were counted and different phenotypes were recorded from one of the absolute methods. Absolute methods were the most similar to one another over time of day and within seasonal dates. Relative methods varied in assessing thrips population dynamics over time of day and within seasonal dates. Based on thrips collected from the plant and sticky card counts, macropterous individuals increased in the spring and summer. Thrips aerially dispersed in the summer. An absolute method, the beat cup method (rapping timothy inside a plastic cup), was among the least variable sampling methods and was faster than direct observations. These findings parallel other studies, documenting the commonality of diel and diurnal effects on sampled arthropod abundance and the seasonal effects on population abundance and structure. These studies also demonstrate that estimated population abundance can be markedly affected by temporal patterns as well as shifting adult phenotypes.     

Cultural and insecticidal control of Thrips tabaci on onions in the Sudan

Onion thrips (Thrips tabaci) is the major pest of onions in the Sudan. In the Gezira Scheme onions are normally transplanted between October and January when the thrips population is very low. Later, thrips commence to breed. Early transplanted onions were well established before infestations became severe and, without the need for insecticidal control, produced significantly higher yields. Onions transplanted later, when thrips began to multiply, produced lower yields. Depending on the duration and intensity of infestation three to five sprays applied at fortnightly intervals were required to control thrips on the late transplanted crop. The effect of spray frequency on the development and yield of onions was investigated and the response found to be solely due to the extent of thrips control achieved. The toxicant itself had no effect on growth. Although the leaves were unaffected by the various treatments, spraying when most thrips were present increased fresh and dry leaf weights, bulb weight and, in the Hudeiba cultivar, the proportion of flowering plants. With the late-transplanted crop, light infestations led to yield losses of at least 39% whilst severe thrips attacks reduced the crop by 57%.     

Euseius addoensis addoensis,an effective predator of citrus thrips,Scirtothrips aurantii,in the eastern Cape Province of South Africa

The predacious mite,Euseius addoensis addoensis (McMurtry) (Acari: Phytoseiidae), significantly (P<0.05) reduced scarring on Palmer Navel oranges,Citrus sinensis (L.) Osbeck, caused by the citrus thrips,Scirtothrips aurantii Faure (Thysanoptera: Thripidae). A chemical exclusion method employing dicofol was used to determine the effect of the predator and numbers ofS. aurantii were monitored on sticky yellow traps. At two sites where numbers of citrus thrips and predacious mites were high, the percentage fruit culled from export quality due to damage byS. aurantii was reduced from 25.5 to 15.5% and from 32.7 to 18.7% by predacious mite activity.Euseius a. addoensis sometimes prevented damage from exceeding 1% cull and was most effective when mean numbers exceeded one mite per inside leaf at petal fall. The best correlation between numbers of mites and the percentage fruit culled at harvest was –0.96 at the location with the highest number of citrus thrips. Tartar emetic (0.398% a.i.) plus sugar (0.4% a.i.) treatments for citrus thrips were slightly detrimental toE. a. addoensis populations but their numbers soon recovered. An alternative treatment, methiocarb 0.016% a.i. plus sugar (0.4% a.i.), almost eliminated the predacious mites and resulted in more citrus thrips damage (10.7% cull) than in the adjacent untreated control (0.2% cull) which contained predacious mites.     

Systemic effects of neem on western flower thrips,Frankliniella occidentalis (Thysanoptera: Thripidae)

The systemic effects of neem on the western flower thrips, Frankliniella occidentalis (Pergande), were investigated in laboratory trials using green bean, Phaseolus vulgaris L., in arena and microcosm experiments. In arena experiments, systemic effects of neem against western flower thrips larvae on primary bean leaves were observed with maximum corrected mortality of 50.6%. In microcosm experiments using bean seedlings, higher efficacy in the control of western flower thrips were observed with soil applications of neem on a substrate mixture (i.e., Fruhstorfer Erde, Type P, and sand) in a 1:1 ratio (93% corrected mortality) compared with application on the commercial substrate only (76% corrected mortality). However, longer persistence of neem was observed with soil application on the commercial substrate, which showed effects against thrips for up to 6 d after application. In addition to systemic effects observed on all foliage-feeding stages of western flower thrips, mortality on contact and repellent effects were observed on soil-inhabiting stages after soil applications of neem. Finally, bean seedlings grown from seeds pregerminated for 3 d in neem emulsion were also toxic to western flower thrips.     

西花蓟马在黄瓜和架豆上的空间分布型及理论抽样数

西花蓟马Frankliniella occidentalis（Pergande）已在北京部分蔬菜园区成功定居,对蔬菜生产造成危害。为了解该虫在蔬菜田的种群空间分布型,指导田间取样,本文应用几种聚集度指标的计算公式以及Taylor、Iwao的回归方程式,分析和测定了西花蓟马的种群空间分布格局。结果表明：在黄瓜和架豆上西花蓟马的空间分布型一致,均为聚集分布型;该虫的空间分布型不受种群密度的影响,并且也不随取样时间的变化而变化。种群数量动态研究显示西花蓟马成虫和若虫在黄瓜植株的中部叶片分布较多（从顶部向下数第4叶至第17叶片上成、若虫的分布数量多于其他叶片上的数量）,未展开的叶片、嫩叶和下部老叶危害较轻。用Iwao回归法中α、β参数计算出在允许误差范围内的理论抽样数。     

Effectiveness of imidacloprid (ProAgro 100 SL) in the control of glasshouse whitefly (Trialeurodes vaporariorum) and western flower thrips (Frankliniella occidentalis) on tomato under cover

In 2002 a glasshouse experiments were carried out on the effectiveness of ProAgro 100 SL in the control of glasshouse whitefly and western flower thrips on tomato cv. Perkoz. Strict observation of the basic prophylactic rules such as introducing in to glasshouse only healthy seedlings, uninfected by whitefly and thrips as well as isolation of tomato plants, especially the seedlings from ornamental plants are important to the limit of the pest population. During the vegetation period the population of whitefly and thrips may be limited by the insecticide application. The glasshouse experiments on the performance of ProAgro 100 SL in the control of mentioned above pest species were carried out at the Research Institute of Vegetable Crops in Skierniewice. The ProAgro 100 SL was applied in the concentration of 0,1%. The standard treatment was Juventox 040 SL (acetamiprid) in the concentration of 0.05%. The obtained results allow to find that ProAgro 100 SL in the rates of 0.1% performed well in the limiting population of whitefly and western flower thrips on glasshouse tomato.     

Monitoring insecticide resistance in Australian Frankliniella occidentalis Pergande (Thysanoptera: Thripidae) detects fipronil and spinosad resistance

Abstract  Insecticide resistance monitoring using a Potter precision spray tower with discriminating concentration and log dose probability techniques underpins the Australian insecticide management strategy for Frankliniella occidentalis Pergande. Abamectin, acephate, chlorpyrifos, dichlorvos, dimethoate, endosulfan, fipronil, malathion, methamidophos methidathion, methiocarb, methomyl, pyrazophos and spinosad are recommended for use against F. occidentalis but abamectin, methiocarb and pyrazophos are the only chemicals where insecticide resistance has not been detected. Although not registered, chlorfenapyr was effective against F. occidentalis and should be pursued for that purpose. In contrast, chlorpyrifos, dichlorvos and malathion resistance were detected at low to moderate levels throughout the study period putting their sustainable use for F. occidentalis control in doubt . Although it appears that acephate, dimethoate, endosulfan, fipronil, methamidophos, methidathion and spinosad remain effective, some populations contained a small percentage of thrips that survived exposure to a concentration that killed 100% of the susceptible strain. Subsequent laboratory selection of one such population separately with fipronil and spinosad caused an increase in resistance to these insecticides. These products must now be considered at risk. This is the first report of fipronil or spinosad resistance in populations of F. occidentalis.