Trichogrammatoidea bactrae Nagaraja (Hymenoptera: Trichogrammatidae) is an important natural enemy of many species of lepidopterous pests. The effects of heat stress temperature (33, 36, and 39 °C), duration of exposure (2, 4, 6, and 8 h), and developmental stage during exposure (embryo-first instar larvae, second instar larvae, prepupae, and pupae) on the development and reproduction of parasitoid T. bactrae were investigated in the laboratory. When exposed to 39 °C for 8 h during pupal stage, only 19.90% adults emerged from host eggs, and more than 14% were deformed (wings were folded or incomplete). Parasitoid females exposed to 39 °C for 8 h as prepupae only lived for 1.45 days and parasitized about 23.5 host eggs. Moreover, life-table parameters of T. bactrae were also influenced by exposure to heat stress temperatures during each preimaginal developmental stage. Based on these results, we propose that T. bactrae is susceptible to high temperatures, especially at 39 °C. Thus, this parasitoid may be more effectively controlling lepidopterous pests during cooler weather conditions.

Nesidiocoris tenuis (Reuter) (Heteroptera: Miridae) is an omnivorous insect used for biological control. Augmentative release and conservation of N. tenuis have been used for pest control in tomato crops. Intracellular bacterial symbionts of arthropods are common in nature and have diverse effects on their hosts; in some cases they can dramatically affect biological control. Fingerprinting methods showed that the symbiotic complex associated with N. tenuis includes Wolbachia and Rickettsia. Rickettsia of N. tenuis was further characterized by sequencing the 16S rRNA and gltA bacterial genes, measuring its amount in different developmental stages of the insect by real-time polymerase chain reaction, and localizing the bacteria in the insect's body by fluorescence in situ hybridization. The Rickettsia in N. tenuis exhibited 99 and 96% similarity of both sequenced genes to Rickettsia bellii and Rickettsia reported from Bemisia tabaci, respectively. The highest amount of Rickettsia was measured in the 5th instar and adult, and the symbionts could be detected in the host gut and ovaries. Although the role played by Rickettsia in the biology of N. tenuis is currently unknown, their high amount in the adults and localization in the gut suggest that they may have a nutritional role in this insect.

Ants act simultaneously as predators and as hemipteran mutualists, and thereby may affect the composition and population dynamics of a wide arthropod community. We conducted ant-exclusion experiments in order to determine the impact of ants on the infestation levels and parasitism of three of the most important citrus pests of western Mediterranean citrus: the honeydew producer Aleurothrixus floccosus Maskell (woolly whitefly) and the non-honeydew producers Aonidiella aurantii Maskell (California red scale; CRS) and Phyllocnistis citrella (Staiton) (citrus leafminer). The study was conducted in three commercial citrus orchards, each one dominated by one ant species (Pheidole pallidula, Lasius grandis or Linepithema humile) during two consecutive growing seasons (2011 and 2012). We registered a significant reduction of the CRS densities on fruits in the ant-excluded treatment in the three orchards and in the two seasons, ranging from as high as 41% to as low as 21%. Similarly, the percentage of shoots occupied by A. floccosus was significantly lower in the ant-excluded plots in the orchards dominated by P. pallidula and L. humile. No significant differences were registered in the percentage of leaf surface loss caused by P. citrella between ant-allowed and ant-excluded treatments in any case. We found no significant differences in the percent parasitism between ant-allowed and ant-excluded treatments for honeydew and non-honeydew producing herbivores. These results suggest that: (i) ant management should be considered in order to reduce herbivore populations in citrus and (ii) mechanisms other than parasitism (e.g., predation) might explain the differences in herbivore infestation levels between treatments.

Plant-mediated interactions between begomoviruses and whiteflies exert important influences on the population dynamics of vectors and the epidemiology of plant diseases. In this article, we synthesize the relevant literature to identify patterns to the interactions. We then review studies on the ecological, biochemical and molecular mechanisms underlying the interactions and finally elaborate on the most interesting issues for future research. The interactions between begomoviruses and the insect vector, the whitefly Bemisia tabaci, via their shared host plants can be mutualistic, neutral or negative. However, in contrast to a pattern of improved performance of vectors on virus-infected plants that has been observed with persistently transmitted RNA viruses, the number of cases exhibiting mutualistic, neutral or negative effects in the indirect interactions between begomoviruses and whiteflies appear evenly distributed. With regard to the mechanisms of plant-mediated positive effects on whiteflies, two case studies indicate that suppression of plant defence and/or alteration in plant nutrition as a result of virus infection can be important. Our review shows that we are only just beginning to understand the tripartite interactions between begomoviruses, whiteflies and plants. Future efforts in this area should try to expand the number and diversity of pathosystems for investigation to reveal the patterns of interactions, to investigate the molecular and biochemical mechanisms of the interactions using a multidisciplinary approach, and to examine the virus–plant–vector interactions in the field and in natural plant communities.

Many hemipteroids are major pests and vectors of microbial pathogens, infecting crops. Saliva of the hemipteroids is critical in enabling them to be voracious feeders on plants, including the economically important ones. A plethora of hemipteroid salivary enzymes is known to inflict stress in plants, either by degrading the plant tissue or by affecting their normal metabolism. Hemipteroids utilize one of the following three strategies of feeding behaviour: salivary sheath feeding, osmotic-pump feeding and cell-rupture feeding. The last strategy also includes several different tactics such as lacerate-and-flush, lacerate-and-sip and macerate-and-flush. Understanding hemipteroid feeding mechanisms is critical, since feeding behaviour directs salivary composition. Saliva of the Heteroptera that are specialized as fruit and seed feeders, includes cell-degrading enzymes, auchenorrhynchan salivary composition also predominantly consists of cell-degrading enzymes such as amylase and protease, whereas that of the Sternorhyncha includes a variety of allelochemical-detoxifying enzymes. Little is known about the salivary composition of the Thysanoptera. Cell-degrading proteins such as amylase, pectinase, cellulase and pectinesterase enable stylet entry into the plant tissue. In contrast, enzymes such as glutathione peroxidase, laccase and trehalase detoxify plant chemicals, enabling the circumvention of plant-defence mechanisms. Salivary enzymes such as M1-zinc metalloprotease and CLIP-domain serine protease as in Acyrthosiphon pisum (Aphididae), and non-enzymatic proteins such as apolipophorin, ficolin-3-like protein and ‘lava-lamp’ protein as in Diuraphis noxia (Aphididae) have the capacity to alter host-plant-defence mechanisms. A majority of the hemipteroids feed on phloem, hence Ca++-binding proteins such as C002 protein, calreticulin-like isoform 1 and calmodulin (critical for preventing sieve-plate occlusion) are increasingly being recognized in hemipteroid–plant interactions. Determination of a staggering variety of proteins shows the complexity of hemipteroid saliva: effector proteins localized in hemipteran saliva suggest a similarity to the physiology of pathogen–plant interactions.

In high-altitude summer pastures, horseflies (Diptera: Tabanidae) can be a serious nuisance to livestock, as well as mechanical vectors of animal diseases such as besnoitiosis, an enzootic disease in the Pyrenees. However, the activity of horseflies in mountainous environments is poorly documented. To study the seasonality and distribution of tabanids in the Pyrenees Mountains, a sampling design was set up in two valleys on opposite sides of the mountain, one north-facing and one south-facing, along high-elevation gradients and at different distances from a water body between May and October 2011. The influence of the landscape on species richness and abundance was assessed by taking into account forested and unforested areas in 200 m radii around the trapping sites. Our findings indicated that: (1) The slope, the altitude and the size of unforested patches significantly influenced community composition of tabanids. (2) Altitude had a positive or a negative effect, depending on the species. (3) Species richness and abundance were negatively correlated with large open habitats and positively correlated with patch-shape complexity. (4) Seasonal succession of the most abundant species was observed in both valleys, with a maximum of catches at the beginning of August; however, tabanid activity ended earlier in the southern valley, which was more exposed to sunlight. (5) Philipomyia aprica, Tabanus bromius, Tabanus glaucopis and Hybomitra auripila were active from 9:00 to 19:00 h (GMT+1), with a peak of activity at midday. This paper also discusses the implications of these findings in relation to changes in horsefly distribution and their control in mountainous environments.

The intensity of pupal diapause in the cotton bollworm, Helicoverpa armigera (Hübner) was investigated under both laboratory and natural conditions. By transferring diapausing pupae induced under LD 11:13, LD 12:12 and LD 13:11 at 20, 22 and 25 °C to 25 °C combined with LD 15:9 to terminate diapause the rearing day length of 11 h evoked greater intensity of diapause than did 12 and 13 h at 25 °C; whereas the rearing temperature of 25 °C evoked more intense diapause than did 20 and 22 °C under LD 11:13. By transferring diapausing pupae induced under LD 12:12 at 20 and 22 °C to six temperatures of 18, 20, 22, 25, 28 and 31 °C combined with LD 15:9 to terminate diapause, the duration of diapause was significantly shortened from 146 days at 18 °C to 24 days at 31 °C, showing that high temperatures significantly accelerate diapause development. Furthermore, the duration of diapause was significantly longer at the rearing temperature of 22 °C than that at 20 °C when the diapause-terminating temperatures were 20 and 22 °C. Chilling at 5 °C did not shorten the duration of diapause but lengthened it when chilling period was included. However, chilling plays an important role in synchronizing adult emergence. Rearing temperature of 22 °C also evoked more intense diapause than did 20 °C in most chilling treatments. When the overwintering pupae were transferred at different times from natural temperatures to 25 °C, it was found that the earlier the transfer took place, the earlier the adults emerged when the time spent under natural conditions was included. However, cool temperatures before March showed an enhanced effect on diapause development at 20 °C, suggesting that the high diapause-terminating temperature can offset the effect of chilling on diapause development. The result of diapause termination under natural conditions suggests that the developmental threshold for post-diapause development in H. armigera should be around 17.5 °C.

We have identified the following three taxa related to the Aphidius colemani species group, which are important biological control agents: Aphidius colemani, Aphidius transcaspicus and Aphidius platensis. Using partial sequences of the mitochondrial cytochrome oxidase subunit I (mtCOI) gene and geometric morphometric analysis of the forewing shape, we have explored the genetic structure and morphological variability of the A. colemani group from different aphid host/plant associations covering a wide distribution area. The topology of the maximum parsimony and maximum likelihood trees were identical with 98–100% bootstrap support, clustering A. colemani, A. platensis and A. transcaspicus into separate species. The distances among the taxa ranged from 2.2 to 4.7%, which is a common rate for the between-species divergence within the subfamily Aphidiinae. Differences in the shape of the forewing investigated within the biotypes of A. colemani group are congruent with their genetic diversification. Both A. platensis and A. colemani share a common host range pattern, and it would be interesting to estimate and compare the role of these two species in future biological control strategies against aphids of economic importance. Our results indicate that ‘genetic screening’ is a reliable approach for identification within the A. colemani group. The high variation in the wing shape among species, including a significant divergence in the wing shape among specimens that emerged from different hosts, makes the forewing shape and wing venation less reliable for species determination. Aphidius platensis is diagnostified and redescribed, and the key for the A. colemani group is presented.

Aphid–plant interactions depend on genotypes of both organisms, which determine the two-way molecular exchange that leads to compatible or incompatible outcomes. The underlying genes are mostly unknown, making it difficult to predict likelihood of aphid success or host resistance, and hampering crop genetic improvement. Here we screened eight pea aphid clonal genotypes collected from diverse legume hosts, on a species-wide panel of Medicago truncatula (Mt) genotypes. Aphid virulence was measured by survival, fecundity and growth rate, together with scores for chlorosis and necrosis as host response indicators. Outcomes were highly dependent on the specific aphid–host genotype combinations. Only one Mt line was fully resistant against all clones. Aphid-induced host chlorosis and necrosis varied greatly, but correlated with resistance only in a few combinations. Bi-clustering analysis indicated that all aphid clones could be distinguished by their performance profiles across the host genotypes tested, with each clone being genetically differentiated and potentially representing a distinct biotype. Clones originating from Medicago sativa ranged from highly virulent to almost completely avirulent on both Medicago species, indicating that some were well adapted, whereas others were most likely migrants. Comparisons of closely related pairs of Australian Mt genotypes differing in aphid resistance revealed no enhanced resistance to European pea aphid clones. Based on the extensive variation in pea aphid adaptation even on unfamiliar hosts, most likely reflecting multiple biotype-specific gene-for-gene interactions, we conclude that robust defences require an arsenal of appropriate resistance genes.

Mutualistic relationships between honeydew-producing insects and ants have been widely recognized for several decades. Iridomyrmex rufoniger (Lowne) is the commonest ant species associated with black scale, Saissetia oleae (Olivier), in the citrus orchards of the mid latitudes of coastal New South Wales. Citrus trees with high densities of both red and black scale and high ant activity were identified and the results of excluding ants from half of those trees (using a polybutene band on each trunk) were compared with the results of not excluding ants from the other half. Trees with a low incidence of black scale and ants were also studied. Exclusion of ants from trees was soon followed by collapse of black scale populations because most individuals were asphyxiated by their own honeydew. Also, parasitism of the red scale by Encarsia perniciosi (Tower) and Encarsia citrina Craw was significantly higher than in the control trees over the following year, as was the predation rate on red scale due to three coccinellid predators, Halmus chalybeus (Boisduval), Rhyzobius hirtellus Crotch and Rhyzobius lophanthae (Blaisdell). In contrast, another coccinellid, Orcus australasiae (Boisduval), and a noctuid moth larva, Mataeomera dubia Butler, were seen in low numbers on banded (ant exclusion) trees, probably because of the low availability of their black scale prey, but were significantly higher on control trees apparently because of their invulnerability to ants.

To understand the effect of plant availability/structure on the population size and dynamics of insects, a specialist herbivore in the presence of two of its parasitoids was studied in four replicated time-series experiments with high and low plant availabilities; under the latter condition, the herbivore suffered from some periods of resource limitation (starvation) and little plant-related structural refuges. Population dynamics of the parasitoid Cotesia vestalis was governed mainly by the delayed density-dependent process under both plant setups. The parasitoid, Diadegma semiclausum, under different plant availabilities and different coexistence situations (either +competitor or –competitor) showed dynamics patterns that were governed mainly by the delayed density process (significant lags at weeks 2–4). Both the competing parasitoids did not experience beneficial or costly interferences from each other in terms of their own population size when the plant resource was limited. Variation in the Plutella xylostella population under limited plant availability is higher than that under the other plant setup. For both parasitoids, under limited plant setup, the extinction risk was lower when parasitoids were engaged in competition, while under the unlimited plant setup, the mentioned risk was higher when parasitoids competed. In this situation, parasitoids suffered from two forces, competition and higher escaped hosts.

The phenology of insect emergence affects reproductive success and is especially critical in short-lived species. An increasing number of studies have documented the effects of thermal and other climatic variations and of unpredictable habitats on the timing of adult insect emergence within and between populations and years. Numerous interacting factors may affect the phenology of adult emergence. Host-plant quality and availability is a key factor that has been largely neglected in studies of the phenology of phytophagous insects. The purpose of this study was to determine the effect of host plant characteristics on the rate of larval growth and the pattern of emergence in a wild population of Lobesia botrana (European grapevine moth), a significant pest in European vineyards. The phenology of emergence differed significantly among the six tested varieties of grapes. The percentage of bunches harboring pupae was similar among the different grape varieties, and the total number of pupae collected was similar to the number of emerging adults per bunch. Among the six varieties of grapes, 0–25 pupae were produced on each bunch. Each of the grape varieties had a single wave of emergence, in which males emerged before females, but their emergence phenology differed significantly in Chardonnay, Chasselas, and Pinot grapes. Both genders had extended durations of emergence in Merlot grapes. Together, the present results show that the characteristics of the grape host plant affect the emergence phenology of L. botrana.

Bactrocera invadens, a fruit fly from Asia, is an invasive pest species across Africa. It appears to continue spreading, not only in latitude but also in altitude. To assess its capacity to infest a large variety of hosts and its competition with other fruit fly species, a study along an altitudinal gradient was conducted. At low altitudes, the high abundance in the field and high infestation of B. invadens in different fruit species make it a serious pest. At high altitudes, colonization has started and B. invadens occurs in low numbers by reproducing successfully in high altitude fruits. Overall the abundance and infestation of B. invadens is influenced by its direct competitor Ceratitis rosa and the presence of its preferred host species. C. rosa is still the dominant species in temperate fruits grown at high altitude. Ceratitis cosyra, however, is negatively affected by B. invadens, this species seems to have shifted hosts to avoid competition. The broad host range and competitive potential of B. invadens increase the risk for further spread not only to higher areas, but also to subtropical regions.

The dispersion of Scaphoideus titanus Ball adults from wild to cultivated grapevines was studied using a novel mark–capture technique. The crowns of wild grapevines located at a distance from vineyards ranging from 5 to 330 m were sprayed with a water solution of either cow milk (marker: casein) or chicken egg whites (marker: albumin) and insects captured in yellow sticky traps placed on the canopy of grapes were analyzed via an indirect ELISA for markers’ identification. Data were subject to exponential regression as a function of distance from wild grapevine, and to spatial interpolation (Inverse Distance Weighted and Kernel interpolation with barriers) using ArcGIS Desktop 10.1 software. The influence of rainfall and time elapsed after marking on markers’ effectiveness, and the different dispersion of males and females were studied with regression analyses. Of a total of 5417 insects analyzed, 43% were positive to egg; whereas 18% of 536 tested resulted marked with milk. No influence of rainfall or time elapsed was observed for egg, whereas milk was affected by time. Males and females showed no difference in dispersal. Marked adults decreased exponentially along with distance from wild grapevine and up to 80% of them were captured within 30 m. However, there was evidence of long-range dispersal up to 330 m. The interpolation maps showed a clear clustering of marked S. titanus close to the treated wild grapevine, and the pathways to the vineyards did not always seem to go along straight lines but mainly along ecological corridors. S. titanus adults are therefore capable of dispersing from wild to cultivated grapevine, and this may affect pest management strategies.

In biological control, successful establishment of a natural enemy species depends on its adaptability in the introduced range including its ability to re-establish desired ecological interactions with the pest. These are affected by genetic parameters hitherto largely unresolved in biological control. The larger grain borer (LGB), Prostephanus truncatus, an invasive species from meso-America, is the most important post-harvest pest of maize in Africa. We studied the genetic structure of Teretrius nigrescens, a predatory beetle previously released for the control of the pest in Africa, to test the hypothesis that establishment patterns were a result of ecotype–environment mismatch and to follow up on our earlier reports of distinct lineages of the predator. We studied 13 populations of T. nigrescens, using 16 polymorphic microsatellite markers. Five genetic populations with a hierarchical structure and significant isolation by distance were detected. The most diverse population was found in southern Mexico, consistent with earlier lineage coexistence observations. Populations introduced to Africa maintained genetic similarity to local geographic populations of their area of origin. The more successful Benin releases were also more genetically diverse. Loss of rare alleles and a higher frequency of existing private alleles in some populations indicated population expansions following bottleneck events. Sustainable biological control should accommodate pest and natural enemy species, and monitor genetic changes associated with introduction and release.

Insect artificial diets are the foundation for mass production of insect predators. Whether there is an influence of long-term rearing with artificial diet on the control ability of predators should be considered. Here, we focused on the effect of nutritional history on the functional response of Geocoris pallidipennis to Myzus persicae. The influence of nutritional history (artificial diet versus natural prey, M. persicae) on the functional response of third to fifth instar nymphs and female G. pallidipennis was examined in the laboratory. The results showed that the functional response curve of both the nymphs and the adult female of G. pallidipennis to M. persicae reflected similar trends on both nutritional histories and confirmed the type II response. Adult female G. pallidipennis reared on either M. persicae or artificial diet produced a significantly better performance than the juvenile stages tested. We estimated that adult female G. pallidipennis can consume 141.6 (artificial diet) or 131.6 (M. persicae) aphids per day, respectively. This indicated that G. pallidipennis reared on both artificial diet and M. persicae displayed high rates of predation.

Lobesia botrana is the most significant pest of grape berries in Spain. Further knowledge of its phenology would enable wine growers to decide on an optimal treatment schedule. The aim of this study is, therefore, to predict the flight peaks of L. botrana in seven wine-growing regions of Spain. The main goal is to provide a prediction model based on meteorological data records. A logistic function model, based on temperature and humidity records, together with an exhaustive statistical analysis, were used to compare the wine-growing regions in which the male flight phenology of L. botrana displays similar patterns and to sort them into groups. By doing so, a joint study of the dynamics of the moth is possible in the regions within each group. A comparison of the prediction errors before and after applying the Touzeau model confirmed that the fit of the latter model is not sufficiently accurate for the regions under study. Moth flight predictions with the logistic function model are good, but accuracy may still be improved by evaluating other non-biotic and biotic factors.

Haemaphysalis longicornis (Acari: Ixodidae) is one of the most common and important arthropod disease vectors in Japan, carrying Japanese spotted fever and bovine theileriosis. The recent expansion of sika deer (Cervus nippon, Artiodactyla: Cervidae) populations, the most common wild host of H. longicornis, has also caused concern about increasing the risk of vector-borne diseases in Japan. We used generalized linear mixed model analysis to determine the relative contribution of deer density and other biological and abiotic factors on the abundance of H. longicornis ticks questing at each developmental stage. A total of 6223 H. longicornis adults, nymphs, and larvae were collected from 70 sites in three regions of central Japan. The abundance of questing adult and nymphal ticks was associated with deer density and other biotic and abiotic factors. However, the abundance of questing larvae showed no association with deer density but did show an association with other biotic and abiotic factors. These findings show that a high density of deer along with other biotic and abiotic factors is associated with increased risk of vector-borne diseases through amplified local abundance of questing nymphal and adult H. longicornis. Further, questing larvae abundance is likely regulated by environmental conditions and is likely correlated with survival potential or the distribution of other host species.

Evolution of resistance to insecticides usually has fitness tradeoffs associated with adaptation to the stress. The basic regulation mechanism of tradeoff between reproduction and resistance evolution to Bacillus thuringiensis (Bt) toxin in the cotton bollworm, Helicoverpa armigera (Ha), based on the vitellogenin (Vg) gene expression was analyzed here. The full-length cDNA of the Vg gene HaVg (JX504706) was cloned and identified. HaVg has 5704 base pairs (bp) with an open reading frame (ORF) of 5265 bp, which encoded 1756 amino acid protein with a predicted molecular mass of 197.28 kDa and a proposed isoelectric point of 8.74. Sequence alignment analysis indicated that the amino acid sequence of HaVg contained all of the conserved domains detected in the Vgs of the other insects and had a high similarity with the Vgs of the Lepidoptera insects, especially Noctuidae. The resistance level to Cry1Ac Bt toxin and relative HaVg mRNA expression levels among the following four groups: Cry1Ac-susceptible strain (96S), Cry1Ac-resistant strain fed on artificial diet with Bt toxin for 135 generations (BtR stands for the Cry1Ac Bt resistance), progeny of the Cry1Ac-resistant strain with a non-Bt-toxin artificial diet for 38 generations (CK1) and the direct descendants of the 135th-generation resistant larvae which were fed on an artificial diet without the Cry1Ac protein (CK2) were analyzed. Compared with the 96S strain, the resistance ratios of the BtR strain, the CK1 strain and the CK2 strain were 2917.15-, 2.15- and 2037.67-fold, respectively. The maximum relative HaVg mRNA expression levels of the BtR strain were approximately 50% less than that of the 96S strain, and the coming of maximum expression was delayed for approximately 4 days. The overall trend of the HaVg mRNA expression levels in the CK1 strain was similar to that in the 96S strain, and the overall trend of the HaVg mRNA expression levels in the CK2 strain was similar to that in the BtR strain. Our results suggest that the changes in reproduction due to the Bt-toxin resistance evolution in the BtR strain may be regulated by the Vg gene expression. The down-regulation of HaVg at the early stages resulted in a period of delayed reproduction and decreased fecundity in the BtR strain. This performance disappeared when the Bt-toxin selection pressure was lost.

The common green lacewing Chrysoperla carnea is a key biological control agent employed in integrated pest management (IPM) programs for managing various insect pests. Spinosad is used for the management of pests in ornamental plants, fruit trees, vegetable and field crops all over the world, including Pakistan. A field-collected population of C. carnea was selected with spinosad and fitness costs and realized heritability were investigated. After selection for five generations, C. carnea developed 12.65- and 73.37-fold resistance to spinosad compared to the field and UNSEL populations. The resistant population had a relative fitness of 1.47, with substantially higher emergence rate of healthy adults, fecundity and hatchability and shorter larval duration, pupal duration, and development time as compared to a susceptible laboratory population. Mean relative growth rate of larvae, intrinsic rate of natural population increase and biotic potential was higher for the spinosad-selected population compared to the susceptible laboratory population. Chrysoperla species are known to show resistance to insecticides which makes the predator compatible with most IPM systems. The realized heritability (h2) value of spinosad resistance was 0.37 in spinosad-selected population of C. carnea.