Costs of insecticide resistance in Cydia pomonella (Lepidoptera: Tortricidae)

The obvious benefits associated with insecticide resistance for pest species may come at a cost to other life-history traits. In this study, we compared the larval and pupal developmental times, pupal mass wing surface area and wing fluctuating asymmetry (FA) in insecticide resistant and control strains of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), collected from apple (Malus spp.) orchards in central Canada. Resistant strains had significantly longer larval developmental times and lower pupal masses compared with the susceptible strain. Although the forewings of resistant moths were smaller in resistant than control strain, no difference in wing FA was detected. Longer developmental times could increase exposure time to natural enemies, and reduced adult size could affect longevity and total reproductive output.     

Genetic analysis of Cydia pomonella (Lepidoptera: Tortricidae) populations with different levels of sensitivity towards the Cydia pomonella granulovirus (CpGV)

Microsatellite (simple sequence repeats, SSR) and mitochondrial DNA markers were used to assess the structure of European codling moth populations showing different levels of susceptibility towards one of the most important biocontrol agents used in apple production, the Cydia pomonella granulovirus CpGV-M. In 638 C. pomonella individuals from 33 different populations a total of 92 different alleles were scored using six SSR loci. The global estimate of genetic differentiation for all 33 populations was not significantly different from zero, thus indicating a lack of genetic differentiation. AMOVA analysis revealed a very weak but significant variance among C. pomonella populations from different geographic regions, however, no significant variation was evident between CpGV-M resistant or susceptible C. pomonella populations. Sequence analysis of a fragment of the cytochrome oxidase subunit 1 in eight C. pomonella populations resulted in 27 haplotypes, which were grouped in two distinct clusters. Again, no genetic differentiation between CpGV-M resistant and susceptible codling moth populations was detectable. In addition, Structure analysis using microsatellites and association tests with mtDNA haplotypes found neither population-level nor individual correlations associated with CpGV-M resistance. Accordingly, this lack of population structure does not allow discriminating between one or several, separate origins of CpGV-M resistance.     

Gene flow among Cydia pomonella (Lepidoptera: Tortricidae) geographic and host populations in South Africa

Information on gene flow among geographic and host populations of C. pomonella (L.) (Lepidoptera: Tortricidae) in South Africa is lacking, despite the importance of these measures for the success of control practices such as chemical control and sterile insect release, which are affected by the amount of gene flow among populations. Therefore, populations collected from nine geographically distant regions in South Africa from apples, pears, and stone fruit were compared using amplified fragment length polymorphism with five selective primer pairs. Results showed that although populations from different hosts were not genetically differentiated, significant evidence for population substructure was apparent between geographic populations. Over local scales, it was possible to distinguish between populations collected from orchards situated <1 km apart. These results suggest that although extensive gene flow occurs among populations from different hosts, gene flow among local geographic C. pomonella populations may be limited and is explained in terms of limited moth flight, the relative isolation of pome fruit production areas, and the absence of wild hosts.     

Trade-off between mobility and fitness in Cydia pomonella L. (Lepidoptera: Tortricidae)

Abstract.  1. The codling moth Cydia pomonella , one of the most important pest insects of apple fruit orchards worldwide, is regarded as a 'sedentary' insect, but the natural populations consist of both mobile and sedentary genotypes that display different dispersal capacity in the field. The current study investigated whether there is a fitness consequence of being mobile in this species by comparing two strains with contrasting levels of mobility obtained through bi-directional selection.
2. Female moths of the sedentary strain were significantly larger, laid more eggs during their lifespan, and both male and female moths of this strain lived longer, when compared with the mobile strain.
3. The age-specific fecundity of female moths from the sedentary strain was significantly greater, in particular between 3 and 5 days after emergence when the peaks of both oviposition and flight capacity occur, relative to the mobile moths, thereby leading to a greater reproductive function.
4. These differences resulted in different demographic parameters between the two strains. The sedentary strain had a significantly greater net reproductive rate ( R _o ) and a higher intrinsic rate of natural increase ( r _m ), although the generation time ( T ) was similar, hence requiring a shorter time to double the population size, in comparison to the mobile strain.
5. This study provides empirical evidence for the cost of being mobile in C. pomonella . Trade-offs between mobility and fitness might have contributed to the existence of genetic variance for dispersal behaviour in the natural populations and the maintenance of relative sedentariness in the species.     

Mating compatibility among populations of codling moth Cydia pomonella Linnaeus (Lepidoptera: Tortricidae) from different geographic origins

The codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is a serious pest of pome fruit worldwide and the sterile insect technique (SIT) provides an environmentally acceptable approach for its control. As the pest is present in both the southern and northern hemispheres it would be possible for a rearing facility in the northern hemisphere to supply sterile moths to an SIT programme in the southern hemisphere during the northern winter and vice versa. This could greatly improve the economics of moth production and the running costs of rearing facilities. However in order to develop this concept, it is important to assess if populations of codling moth from different geographical regions share mating compatibility. Twelve different laboratory and field populations from both hemispheres were sampled and field cage bisexual mating compatibility tests were carried out between selected combinations. The index of sexual isolation (ISI) and the female and male relative performance index (FRPI and MRPI, respectively) were calculated for each mating combination. In only two of the combinations was there a slight but significant deviation from random mating. There were also some significant differences in mating duration between the homotypic matings and the duration of a particular homotypic mating seemed to depend on the origin of the other population in the cage. It was concluded that there exist no barriers to mating between populations of codling moth from many parts of the world and that it would be feasible for sterile moths to be shipped from one rearing facility to SIT programmes in other parts of the world.     

Effects of chlorpyrifos on enzymatic systems of Cydia pomonella (Lepidoptera: Tortricidae) adults

The control program of codling moth (Cydia pomonella L.) in the Río Negro and Neuquén Valley is intended to neonate larvae. However, adults may be subjected to sublethal pesticide concentrations generating stress which might enhance both mutation rates and activity of the detoxification system. This study assessed the exposure effects of chlorpyrifos on target enzyme and, both detoxifying and antioxidant systems of surviving adults from both a laboratory susceptible strain (LSS) and a field population (FP). The results showed that the FP was as susceptible to chlorpyrifos as the LSS and, both exhibited a similar chlorpyrifos‐inhibitory concentration 50 (IC₅₀) of acetylcholinesterase (AChE). The FP displayed higher carboxylesterase (CarE) and 7‐ethoxycoumarine O‐deethylase (ECOD) activities than LSS. Both LSS and FP showed an increase on CarE activity after the exposure to low‐chlorpyrifos concentrations, followed by enzyme inhibition at higher concentrations. There were no significant differences neither in the activities of glutathione S‐transferases (GST), catalase (CAT) and superoxide dismutase (SOD) nor in the reduced glutathione (GSH) content between LSS and FP. Moreover, these enzymes were unaffected by chlorpyrifos. In conclusion, control adults from the FP exhibited higher CarE and ECOD activities than control adults from the LSS. AChE and CarE activities were the most affected by chlorpyrifos. Control strategies used for C. pomonella, such as rotations of insecticides with different modes of action, will probably delay the evolution of insecticide resistance in FPs from the study area.     

Isolation of microsatellite loci in the codling moth,Cydia pomonella (Lepidoptera: Tortricidae)

Cydia pomonella L. is an important insect pest of pome fruits worldwide. We have isolated and characterized 17 microsatellite loci from the enriched genomic libraries constructed using a biotin/streptavidin capture protocol. Among these loci, 11 were scored polymorphic when 50 individuals from a laboratory population were screened. Their alleles numbered two to four, with the observed heterozygosity (H_O) ranging from 0.114 to 0.404. Successful amplification was obtained for all these loci when the designed primers were tested, showing the promise of use in genetic mapping and population studies.     

Rapid thermal responses and thermal tolerance in adult codling moth Cydia pomonella (Lepidoptera: Tortricidae)

In order to preserve key activities or improve survival, insects facing variable and unfavourable thermal environments may employ physiological adjustments on a daily basis. Here, we investigate the survival of laboratory-reared adult Cydia pomonella at high or low temperatures and their responses to pre-treatments at sub-lethal temperatures over short time-scales. We also determined critical thermal limits (CTLs) of activity of C. pomonella and the effect of different rates of cooling or heating on CTLs to complement the survival assays. Temperature and duration of exposure significantly affected adult C. pomonella survival with more extreme temperatures and/or longer durations proving to be more lethal. Lethal temperatures, explored between −20 °C to −5 °C and 32 °C to 47 °C over 0.5, 1, 2, 3 and 4 h exposures, for 50% of the population of adult C. pomonella were −12 °C for 2 h and 44 °C for 2 h. Investigation of rapid thermal responses (i.e. hardening) found limited low temperature responses but more pronounced high temperature responses. For example, C. pomonella pre-treated for 2 h at 5 °C improved survival at −9 °C for 2 h from 50% to 90% (p < 0.001). At high temperatures, pre-treatment at 37 °C for 1 h markedly improved survival at 43 °C for 2 h from 20% to 90% (p < 0.0001). We also examined cross-tolerance of thermal stressors. Here, low temperature pre-treatments did not improve high temperature survival, while high temperature pre-treatment (37 °C for 1 h) significantly improved low temperature survival (−9 °C for 2 h). Inducible cross-tolerance implicates a heat shock protein response. Critical thermal minima (CTmin) were not significantly affected by cooling at rates of 0.06, 0.12 and 0.25 °C min⁻¹ (CTmin range: 0.3-1.3 °C). By contrast, critical thermal maxima (CTmax) were significantly affected by heating at these rates and ranged from 42.5 to 44.9 °C. In sum, these results suggest pronounced plasticity of acute high temperature tolerance in adult C. pomonella, but limited acute low temperature responses. We discuss these results in the context of local agroecosystem microclimate recordings. These responses are significant to pest control programmes presently underway and have implications for understanding the evolution of thermal tolerance in these and other insects.     

Attract and kill for codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) control in Syria

The possibility of controlling the codling moth Cydia pomonella (Linnaeus) using an attract and kill approach as an alternative to chemical sprays with contact insecticides was investigated in widely separated orchards. The results of a 4‐year study have shown that, using an attract and kill approach, three applications/season kept infestation rates in treated orchards below the economic injury level except in one with a too high codling moth population density. The mean number of male codling moths/trap/week in attract and kill‐treated orchards was much lower in comparison with control orchards which were treated with the usual cover sprays of insecticides. The results also showed that the efficacy of attract and kill under orchard conditions decreased with time and the relationship between time effect and codling moth death rate was very strong. These data indicate that the attract and kill technique applied at a rate of three application per season resulted in good control of codling moth in well managed orchards in Syria.     

Effects of carbaryl and cyhexatin on survival and reproductive behaviour of Cydia pomonella (Lepidoptera: Tortricidae)

The effects of carbaryl, an insecticide and an abscission agent, and cyhexatin, an acaricide, on the survival, reproductive behaviour and subsequent egg hatch and larval migration of the codling moth, Cydia pomonella L., were investigated under laboratory conditions. Dilution series starting with the concentration recommended for high volume application in the field were evenly sprayed in plastic cups. Depending on the concentration used, both pesticides resulted in high adult mortality, males being more susceptible. Although the total number of the eggs laid by C. pomonella females was significantly reduced in most treatments, the females exposed to different residues of carbaryl laid significantly more eggs during the first 24 h of exposure. This was not the case for cyhexatin. While in all treatments almost 50% of the eggs were laid on the side of the cups, the spatial distribution of the eggs laid under the lid and on the floor was concentration dependent. With one exception, continuous exposure of the moths to carbaryl significantly reduced the number of spermatophores per female. Less than one spermatophore per female was transferred at the highest four concentrations of carbaryl and the highest three concentrations of cyhexatin; whereas, the number of spermatophores per female amounted to about four per female for the other treatments. In contrast to carbaryl, which exhibited a high ovicidal effect at all concentrations, the ovicidal activity of cyhexatin was low. This was manifested in a high rate of egg hatch and the extent of embryonic development in most of the treatments. Furthermore, while carbaryl killed the newly-hatched larvae at all concentrations, cyhexatin acted as a larvicide at the highest two concentrations only.     

苹果蠹蛾滞育特性研究进展

苹果蠹蛾是苹果、梨等仁果类果树上的重要害虫,也是重要的入侵害虫。其具有寄主广泛、适生性强、蛀果为害等特点,一旦侵入很难彻底根除。老熟幼虫脱果后寻找隐蔽场所滞育的习性是其适应不利环境和维持种群繁衍的重要生存策略。本文就苹果蠹蛾幼虫滞育的诱导和终止因素、滞育场所的选取及聚集习性、滞育幼虫对环境的适应能力及内在生理生化特性以及针对滞育幼虫的防治技术等研究进行了综述,为探明苹果蠹蛾的灾变机理、预测预报和防治提供依据。     

Mating behaviors of Cydia pomonella (Lepidoptera: Tortricidae) as influenced by sex pheromone in electrostatic powder

Entostat is an electrostatically charged wax powder that can adhere strongly to insect cuticle, making it an ideal carrier to deliver pheromone for pheromone-based confusion techniques. We investigated the attractiveness of Cydia pomonella (L.) (Lepidoptera: Tortricidae) males treated with pheromone-laden Entostat powder to naive conspecifics as well as mating behaviors of males after such treatment in a laboratory flight tunnel. Male moths exposed to Entostat containing 1% E,E-8,10-dodecadien-1-ol (codlemone) acquired and retained the largest amount of the powder and became more attractive as point sources to naive moths compared with those treated with powder containing 5 or 10% codlemone. All Entostat-exposed males remained as attractive as a 0.1-mg codlemone lure for up to 24 h in flight tunnel investigations. Male moth orientation to normally attractive sources of codlemone was completely disrupted directly after treatment with Entostat powder. Males' ability to orient to 0.1-mg lures recovered progressively over a 6-d postexposure interval; however, their responses never reached the levels observed with untreated control moths. Entostat-exposed moths retained detectable amounts of codlemone up to 4 d. Our laboratory flight tunnel results suggest that the mode of action of Entostat powder as an autodissemination control tactic may be due to creating both attractive false point sources after exposure to the powder as well as directly inhibiting contaminated males' capability to orient to pheromone.     

Thermal conditioning of fifth-instar Cydia pomonella (Lepidoptera: Tortricidae) affects HSP70 accumulation and insect mortality

Abstract.  Levels of HSP70 protein of fifth-instar codling moth [ Cydia pomonella (L.) (Lepidoptera: Tortricidae)] are determined after conditioning at 35 °C for different times and also after recovery at 22 °C. Protein samples from larvae conditioned for different times are separated by sodium dodecyl sulphate-polyacrylamide gel electrophoresis electrophoresis. Sub-lethal thermal conditioning at 35 °C for 40 min, 2, 6 and 18 h induces new protein bands in the extracts from treated codling moth larvae. Immunodetection with an antibody to a heat-inducible HSP70 indicates a stronger reaction after 35 °C for 2, 6 and 18 h than after 35 °C for 40 min or control and, during the recovery period at 22 °C, the level of heat shock protein decreases. Conditioning of fifth-instar codling moths at 35 °C also induces thermotolerance in the insects and necessitates longer times at a lethal temperature to ensure mortality. Thermotolerance is correlated with the accumulation of heat inducible HSP70 protein.     

Developing female removal for Cydia pomonella (Lepidoptera: Tortricidae) in organic pear in the USA and Italy

Seventeen trials were conducted using traps baited with kairomone-based lures to evaluate female removal (FR) as an effective management tactic for codling moth, Cydia pomonella, L., in organic pear, Pyrus communis L., production in Italy and USA during 2019–2020. Studies included paired plots (0.4–1.4 ha) treated with or without 60 traps ha⁻¹ in cultivars Bartlett and Abate Fétel. Paired plots were also treated together with or without mating disruption (MD) and with similar spray programmes. Three-, four-, and five-component lures were used with several trap types: green, clear or green/white bucket traps and orange delta traps. The three-component lure consisted of (E,E)-8,10-dodecadien-1-ol (sex pheromone, PH), (E,Z)-2,4-ethyl decadienoate (pear ester, PE) and acetic acid (AA); the four-component lure was PE, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), 6-ethenyl-2,2,6-trimethyloxan-3-ol (pyranoid linalool oxide, LOX), and AA, and the five-component lure had PH added. Preliminary studies were also conducted to evaluate the trapping efficacy of non-saturating bucket traps. A clear bucket trap baited with PE/DMNT/LOX + AA captured 97-fold more codling moth females than an orange delta trap baited with PH/PE. However, female captures did not differ between traps when both were baited with the four-component lure. Fruit injury from codling moth was significantly reduced with the implementation of FR in both pear production regions. At harvest, mean fruit protection in the USA studies was 65% and 27% in Italy. This difference was likely associated with the four-component lure being less effective in Italy than in the USA. Nevertheless, results demonstrate that FR can be a useful approach to remove females immigrating into orchards and as a new tactic to reduce pest pressure in selected areas of orchards allowing both MD and organic insecticide programmes to be more effective. Further studies should investigate the cost-benefit of matching the intensity of FR to variable pest pressures.     

Fruit Bins Washed with Steinernema carpocapsae (Rhabditida: Steinernematidae) to Control Cydia pomonella (Lepidoptera: Tortricidae)

A bin washing apparatus (drencher) was used to treat late-instar codling moth, Cydia pomonella Linnaeus-infested wooden fruit bins with the entomopathogenic nematode Steinernema carpocapsae . Sentinel late-instar codling moth larvae were placed in strips of wood, made from dismantled bins, that were subsequently screwed into the corners of real bins before treatment. Pre-soaking the bins with water prior to treatment and covering the bins with a plastic tarp post-treatment, to maintain a high humidity, significantly increased sentinel codling moth mortality. Treatment for 1 min with 50 S. carpocapsae infective juveniles mL -1 water resulted in more than 80% mortality over 6 h in which the nematodes were continuously pumped through the drencher. The temperature of the water in the bin washer was ≤15°C and the chlorine in the city water was too low to harm the nematodes. The key disadvantage of washing the bins was the labour involved. It may be possible to reduce post-treatment holding time and increase codling moth mortality by holding the bins at a higher temperature post-treatment, however this parameter was not included in the study. The presence of the codling moth granulosis virus within the larvae did not influence the nematode-induced codling moth mortality over a range of concentrations and two temperatures in laboratory trials.     

Resistance and cross-resistance to four insecticides in populations of obliquebanded leafroller (Lepidoptera: Tortricidae)

Populations of obliquebanded leafroller, Choristoneura rosaceana (Harris), were collected from organic and conventionally managed orchards located in the Okanagan and Similkameen Valleys of British Columbia Neonate F1 progeny were assayed for resistance to azinphosmethyl, tebufenozide, methoxyfenozide, and indoxacarb using a leaf disk bioassay. Significant differences in resistance levels among populations were observed for all four insecticides. Insects collected from organic sites were more susceptible to all insecticides than were insects collected from conventional sites. Resistance to the benzoylhydrazine insect growth regulators tebufenozide and methoxyfenozide was highly correlated with resistance to azinphosmethyl across populations, indicating cross-resistance between these compounds. The highest levels of resistance were observed with indoxacarb, but resistance levels to indoxacarb did not correlate with those for azinphosmethyl. Dose-response regression lines for tebufenozide were parallel across populations, suggesting that the resistance mechanism(s) were quantitatively, but not qualitatively, different. Cross-resistance between azinphosmethyl and benzoylhydrazine insecticides indicates that a resistance management strategy for obliquebanded leafroller involving the rotation of these materials is not likely to be successful.     

Control of Cydia pomonella L. and Cydia molesta (Busck) (Lepidoptera Tortricidae) in pome-fruit orchards with Ecodian sex pheromone dispensers

A mating disruption approach using high densities of pheromone dispensers, has been recently proposed for codling moth, Cydia pomonella (L.), and oriental fruit moth, Cydia molesta (Busck.), (Lepidoptera Tortricidae), control. Ecodian Star dispensers, made of low-cost biodegradable material and easy to apply, were formulated with 10 mg of codlemone (E8,E10-12OH) and 10 mg of grapamone (Z8-12OH) and placed at a rate of 1,400-2,000 dispensers/ha. The pheromone release rates from new and field aged dispensers were evaluated by hexane extraction of the residual attractant (indirectly) and gas-chromatographic analysis. The release rate of field-aged dispensers decreased over time with a good linearity; they released a significant amount of synthetic sex pheromones over the entire season. Dispensers elicited close-range approaches of codling moth males in wind tunnel irrespective of their age. Field trials carried out from 2003 to 2004 confirmed the efficacy of Ecodian Star dispensers for codling moth and oriental fruit moth control, regardless the size of the treated area. Our results demonstrate that Ecodian dispensers achieved a good level of activity and longevity over the season. The potential of this strategy for the control of the moths is discussed.     

Rapid assessment of the sex of codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) eggs and larvae

Two different methods were tested to identify the sex of the early developmental stages of the codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) with a WZ/ZZ (female/male) sex chromosome system. First, it was shown that the sex of all larval stages can be easily determined by the presence or absence of sex chromatin, which is formed by the female‐specific W chromosome in interphase nuclei. This trait can also be used to identify the sex of newly hatched larvae but it does require care and accuracy. Secondly, a new sexing technique was developed based on a molecular marker of the codling moth W chromosome. Flanking regions of an earlier described W‐specific sequence (CpW2) were isolated and sequenced and a 2.74 kb sequence (CpW2‐EcoRI), specific for the W chromosome, was obtained. Several PCR tests were conducted, which confirmed that the CpW2‐EcoRI sequence is a reliable marker for the sex identification in codling moth samples of different geographical origin. In addition, a fragment of a codling moth gene, period (Cpper) was isolated and sequenced. Results of southern hybridization of the Cpper probe with female and male genomic DNA suggested that the Cpper gene is located on the Z chromosome. Then a multiplex PCR assay was developed, which co‐amplified the CpW2‐EcoRI sequence to identify the W chromosome and the Z‐linked Cpper sequence, which served as a positive control of accurate processing of tested samples. The multiplex PCR provides an easy and rapid identification of the sex of embryos and early larval instars of the codling moth.