Effect of rearing strategy and gamma radiation on fecundity and fertility of codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae)

The codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is a serious pest of pome fruit worldwide. In an effort to reduce the use of pesticides to control this pest, the sterile insect technique (SIT) is used or considered for use as a component of area‐wide integrated pest management programmes. Rearing codling moths through diapause has been shown to improve the competitiveness of sterile moths released in orchards, and provides management alternatives that would allow mass‐rearing facilities to increase their yearly production of sterile moths. Because radiosensitivity in insects can be influenced by numerous biological factors, laboratory tests were conducted to examine whether the response to increasing doses of radiation, as expressed in the fecundity and fertility of cohorts of moths, is similar for adult moths mass‐reared through diapause or through standard (non‐diapause) production protocols. Our data revealed that the effect of increasing doses of radiation on fecundity and fertility of codling moths reared through both rearing strategies was similar. In the case of fertility, this is a particularly important finding for the expanded application of codling moth SIT. If mass‐rearing facilities use year‐round diapause rearing, the dose required to treat the insects prior to release will be similar to that used when codling moths are reared through standard production protocols.     

Climate change effects on walnut pests in California

Increasing temperatures are likely to impact ectothermic pests of fruits and nuts. This paper aims to assess changes to pest pressure in California's US$0.7 billion walnut industry due to recent historic and projected future temperature changes. For two past (1950 and 2000) and 18 future climate scenarios (2041–2060 and 2080–2099; each for three General Circulation Models and three greenhouse gas emissions scenarios), 100 years of hourly temperature were generated for 205 locations. Degree‐day models were used to project mean generation numbers for codling moth (Cydia pomonella L.), navel orangeworm (Amyelois transitella Walker), two‐spotted spider mite (Tetranychus urticae Koch), and European red mite (Panonychus ulmi Koch). In the Central Valley, the number of codling moth generations predicted for degree days accumulated between April 1 and October 1 rose from 2–4 in 1950 to 3–5 among all future scenarios. Generation numbers increased from 10–18 to 14–24 for two‐spotted spider mite, from 9–14 to 14–20 for European red mite, and from 2–4 to up to 5 for navel orangeworm. Overall pest pressure can thus be expected to increase substantially. Our study did not include the possibility of higher winter survival rates, leading to higher initial pest counts in spring, or of extended pest development times in the summer, factors that are likely to exacerbate future pest pressure. On the other hand, initiation of diapause may prevent an extension of the season length for arthropods, and higher incidence of heat death in summer may constrain pest population sizes. More information on the impact of climate change on complex agroecological food webs and on the response of pests to high temperatures is needed for improving the reliability of projections.     

Regulation of heat shock proteins in the apple maggot Rhagoletis pomonella during hot summer days and overwintering diapause

Abstract Developing larvae of the apple maggot Rhagoletis pomonella are frequently exposed to summertime apple temperatures that exceed 40 °C and, during their overwintering diapause, pupae are exposed to sub‐zero soil temperatures for prolonged periods. To investigate the potential involvement of heat shock proteins (Hsps) in response to these environmental extremes, the genes encoding Hsp70 and Hsp90 in R. pomonella are cloned and expression monitored during larval feeding within the apple and during overwintering pupal diapause. Larvae reared in the laboratory at constant temperatures of 25, 28 or 35 °C express Hsp90 but very little Hsp70. Larvae do not survive rearing at 40 °C. The temperature cycles to which larvae were exposed inside apples in the field, ranging 16–46.9 °C over a 24‐h period, elicit strong Hsp70 and Hsp90 expression, which begins at mid‐day and reaches a peak in late afternoon, coinciding with peak air and apple temperatures. Heat shock proteins are also expressed strongly by pupae during their overwintering diapause. Hsp70 is not expressed in nondiapausing pupae but is highly expressed throughout diapause. Hsp90 is constitutively expressed in both diapausing and nondiapausing pupae. Rhagoletis pomonella thus strongly expresses its Hsps during pupal diapause, presumably as a protection against low temperature injury, and during larval development to cope with natural temperature cycles prevailing in late summer.     

Predicting the phenology of codling moth,Cydia pomonella,for sustainable pest management in Swiss apple orchards

The effects of temperature on post‐diapause development, female lifespan, and reproduction of Cydia pomonella L. (Lepidoptera: Tortricidae) were studied under controlled conditions to determine the respective parameters for a temperature‐driven phenology model. Lower thermal thresholds of 10.0 and 9.7 °C and thermal constants of 417.2 and 427.7 degree days were established for post‐diapause development of females and males, respectively. Female lifespan was found to be 202.6 degree days on average, with a lower thermal threshold of 11.2 °C. Total fecundity was highest at 27.7 °C with an average of 241.9 eggs per female. Literature data were used to quantify the lower thermal thresholds and the thermal constants for eggs, larvae, and pupae. For validation, the model was run with weather data from 13 consecutive years and the output was compared with pheromone trap catches from 39 to 63 different orchards, depending on the year. The model proved to be a reliable and useful tool to forecast codling moth phenology under Swiss conditions not only for a whole growing season but also during several years with very variable weather conditions. Hence, it can be used to support apple growers in pest management decisions on the one hand and to predict changes in codling moth phenology with respect to climate change on the other.     

The diapause response of Rhagoletis pomonella to varying environmental conditions and its significance for geographic and host plant‐related adaptation

The recent shift of Rhagoletis pomonella Walsh (Diptera: Tephritidae) from its ancestral host hawthorn to apple is a model for incipient sympatric speciation in action. Previous studies have shown that changes in the over‐wintering pupal diapause are critical for differentially adapting R. pomonella flies to a difference in the fruiting times of apples vs. hawthorns, generating ecologically based reproductive isolation. Here, we exposed pupae of the hawthorn race to various combinations of pre‐ and over‐wintering rearing conditions and analyzed their effects on eclosion time and genetics. We report certain unexpected results in regards to a combination of brief pre‐winter and over‐wintering periods indicative of gene*environment interactions requiring a reassessment of our current understanding of R. pomonella diapause. We present a hypothesis that involves physiological factors related to stored energy reserves in pupae that influences the depth and duration of Rhagoletis diapause. This ‘pupal energy reserve’ hypothesis can account for our findings and help clarify the role host plant‐related life history adaptation plays in phytophage biodiversity.     

Population genetic structure of Cydia pomonella: a review and case study comparing spatiotemporal variation

Analysis of population genetic structure is a key aspect to understand insect pest population dynamics in agricultural scenarios. Here the role of geography, hosts and time on the population genetic structure of codling moth Cydia pomonella (Linnaeus) (Lep., Tortricidae) populations is described. Temporal variation was examined in two French orchards among each of three adult flights during two successive years. Analyses were conducted using two insecticide resistance markers (variation at the sodium channel gene and enzymatic activity of cytochrome P450 oxidases) and three microsatellite loci. Levels of genetic variation among temporal populations were not significant based on variation in the sodium channel gene and microsatellite loci. However, P450 oxidase activity differed significantly during both flights and years, decreasing during the three flights of the first year and increasing during the second. These results suggest that phytosanitary measures are among the factors shaping the genetic structure of C. pomonella populations over temporal and geographical scales. We discuss the relative importance of natural and passive dispersal related to anthropogenic activities affecting C. pomonella population genetics and highlight population genetic research needs in order to design more efficient pest management practices.     

Testing temperature effects on woodboring beetles associated with oak dieback

A warmer climate may potentially have a strong effect on the health status of European oak forests by weakening oak trees and facilitating mass reproduction of wood boring insects. We did a laboratory experiment in Slovakia to study the response of major pest beetles of oak and their parasitoids to different temperature regimes as background for predicting climatic effects and improving management tools of European oak forests. With higher temperatures the most important oak pest Scolytus intricatus emerged much earlier, which indicate that completion of a second generation and increased damage further north in European oak forests may be possible. Lower temperatures gave longer larval galleries and more offspring per parents but still lower beetle production due to semivoltine life cycle. For buprestids and longhorn beetles warmer temperatures resulted in more emerging offspring and a shift towards earlier emergence in the same season, but no emergence in the first season indicated that a change to univoltine populations is not likely. Reduced development success of parasitoids at the highest temperatures (25/30 °C) indicates a loss of population regulation for pest beetle populations. A warmer climate may lead to invasion of other population-regulating parasitoids, but also new serious pest may invade. With expected temperature increases it is recommended to use trap trees both in April and in June, and trap trees should be removed within 2 months instead 1 year as described in the current standard.     

Spatial and temporal distribution of bioclimatic potential for the Codling moth and the Colorado potato beetle in Norway: model predictions versus climate and field data from the 1990s

Abstract 1 Based on climate data from a network of agrometeorological stations in Norway, the effects of current and future climate regimes on the spatial and temporal distribution of the Codling moth (Cydia pomonella) and the establishment potential of the Colorado potato beetle (Leptinotarsa decemlineata) were investigated. 2 The study was accomplished using climex , a dynamic climate matching‐ and climate response estimation model, which predicts potential distribution of an organism based on its known geographical distribution. 3 Validation of the climex model predictions for C. pomonella against field data on spatial distribution of the species in Norway resulted in a refined set of climate response parameters for C. pomonella. Temporal occurrence of C. pomonella seems to be affected by climate (temperature) and insecticide treatment against the Apple fruit moth (Argyresthia conjugella) in the previous season. 4 Climate change scenarios (0.1 °C increase per degree in latitude in daily maximum and minimum temperatures) indicated an extension of the potential geographical range for C. pomonella, and 23 new locations were found favourable for its long‐term survival. The abundance and pest status of C. pomonella could increase dramatically in those locations where the species is already established. 5 Leptinotarsa decemlineata would only temporarily find suitable climate conditions in Norway and hence only be able to establish interim populations in a few regions under current climate conditions. Climate change scenarios for L. decemlineata indicated that the species would be able to establish as far north as 64°N, mainly in the inland of eastern Norway. 6 In general, the methods applied support the process of decreasing the uncertainty both in our knowledge about the pests themselves and about the environment, which are crucial elements in predicting whether a species is able to establish in a new area.     

Diapause induction in the codling moth, Cydia pomonella: effect of prediapause temperatures

The effect of four prediapause temperatures (18, 22, 26 and 30°C) on the photoperiodic response of the codling moth, Cydia pomonella (L.), was studied under controlled conditions. The highest rates of diapause were recorded, for all day-lengths, at temperatures of 22 and 26°C while relatively lower rates of diapause were elicited at 18 and 30°C. The same trend was demonstrated by projecting the values of the critical photoperiod which induces 50% diapause (=CPhP₅₀) over the prediapause temperature. The change in diapause incidence as a function of photoperiod, at all prediapause temperatures, exhibited a response characteristic of long-day insects, i.e. high rates of diapause at short days (12–13.5 h) and a decrease in diapause incidence at long days (14–15 h). The results for temperatures 22, 26 and 30°C support the view that lower prediapause temperatures enhance diapause induction, at a give photoperiod, while higher temperatures tend to avert or diminish the process. On the other hand, the low rates of diapause obtained at 18°C contradict this view. Nevertheless, high correlation was found between the laboratory evidence and field data, indicating the adaptability of the Israeli codling moth to subtropical climate.     

Climatic limits of pink bollworm in Arizona and California: effects of climate warming

The distribution and abundance of pink bollworm (Pectinophora gossypiella Saunders (PBW)) in cotton in Arizona and California was examined using a validated weather-driven, physiologically based demographic model of cotton and PBW integrated into a geographic information system (GIS). Survival of diapause larvae during winter as affected by low temperatures is a key factor determining the range of PBW. Winter survival was estimated using data from Gutierrez et al. (Can. Entomol. 109 (1977) 1457) and Venette et al. (Environ. Entomol. 29 (5) (2000) 1018). The model was run continuously over the period 1 January 1995 to 31 December 2003 using observed weather data from 121 locations. Three output variables were mapped as measures of PBW persistence: over-winter survival of diapause PBW larvae, cumulative daily PBW larval densities over the season, and the number of diapause larvae produced during the season. The distribution of pink bollworm is predicted to be restricted to the relatively frost-free cotton growing areas of Arizona and Southern California where it currently reaches pest status. The model predicts that extension of PBW's range into the Central Valley of California is unlikely. The analysis questions the efficacy of an ongoing area-wide effort to prevent the establishment of PBW in the Central Valley of California. Four global warming scenarios were examined to estimate the effects on the potential geographic range of PBW. Average observed daily temperatures were increased 1.0, 1.5, 2.0 or 2.5 °C, respectively, in the four scenarios. Scenarios with average increases of 1.5–2.5 °C predicted that the range of PBW would expand into the Central Valley of California and the severity of the pest would greatly increase in areas of current infestation.     

Diapause termination of Rhagoletis cerasi pupae is regulated by local adaptation and phenotypic plasticity: escape in time through bet‐hedging strategies

Persistence and thriving of univoltine, herbivore insect species of the temperate zone rely on obligate diapause response that ensures winter survival and synchronization with host phenology. We used a stenophagous fruit fly (Rhagoletis cerasi) with obligate pupae diapause to determine genetic and environmental effects on diapause intensity of geographically isolated populations with habitat heterogeneity. Pupae from two Greek and one German populations with various gene flow rates were exposed at five constant chilling temperatures (0–12 °C) for different durations and then incubated at a high temperature until all adults have emerged. Pupae diapause intensity differs among Greek and German populations, suggesting an adaptive response to habitat heterogeneity (mostly differences in phenology patterns of local host cultivars). Moderately warm winter temperatures, such as 8 °C, promote diapause termination in all three populations. Insufficient chilling (short duration or warmer temperatures) regulates the expression of prolonged dormancy. Interestingly, extended chilling (longer than required for terminating diapause) ‘return’ pupae to another (facultative) cycle of dormancy enabling adults to emerge during the next appropriate ‘window of time’; a strategy first time reported for univoltine insects. Consequently, diapause duration of R. cerasi is determined both by i) the adaptive response to local climatic conditions (annual dormancy) and ii) the plastic responses to interannual climatic variability resulting in two types of long life cycles within populations, prolonged and facultative dormancy as response to insufficient chilling and extended exposure to chilling, respectively. Long life cycles are expressed as a part of dormancy bet‐hedging strategies of R. cerasi populations.     

Comparison of fatty acid composition in total lipid of diapause and non-diapause larvae of Cydia pomonella （Lepidoptera： Tortricidae）

Seasonal changes in the fatty acid composition of the total lipid extracted from the whole body of Cydia pomonella L. larvae were determined by gas chromatography. The six most abundant fatty acids in both non-diapause and diapause larvae of codling moth were oleic （35%-39%）, palmitic （23%-33%）, linoleic （16%-30%）, palmitoleic （5%-10%）, stearic （1.5%-3.0%） and linolenic acids （1.0%-2.5%）. This represents a typical complement of Lepidopteran fatty acids. The fatty acid composition of total lipid of C. pomonella larvae was related to diapause. In similarity to most other reports, the proportion of unsaturated fatty acids increased in diapause initiation state. The total lipid of diapause larvae contained more linoleic acid （25.8% vs. 16.1%） and less palmitic acid （24.7% vs. 33.4%）, than that of non-diapause larvae. The weight percentage of linoleic acid （C 18：2） increased from 16% to 26% from early-August through early-September during transition to diapause, while palmitic acid （C16：0） decreased from 33% to 25% at the same time. These changes resulted in an increase in the ratio of unsaturated to saturated fatty acids （UFA/SFA） from 1.72 in non-diapause larvae to 2.63 in diapause larvae.     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Diapause development in early and late emerging phenotypes of Delia floralis

The turnip fly, Delia floralis Fall6n （Diptera： Anthomyiidae） is an important insect pest of brassica vegetable crops in the holarctic region. Different populations have strongly varying temperature requirements for fly emergence, a challenge for accurate prediction of activity. This study focused on diapause development in one early and one late emerging phenotype. The physiological state after various treatments was deduced from emergence data. Our results showed a slow diapause progression at chilling conditions for both populations and diapause ended about 7 months after pupae were formed for the early population. For the late population held at 4℃ diapause did not end, no matter how long the duration of chilling. These pupae required a period with elevated temperatures above 6~C to continue development. At constant non-chilling conditions （18℃） from the time pupae were formed both populations completed diapause most rapidly. These results indicate that chilling delayed, rather than accelerated development and was not a prerequisite for diapause development. For post-diapause, results indicated a linear relationship between rate of development and temperature within the range of 6-18℃and a theoretical base temperature for development of about 2℃ for both populations. In conclusion, D. floralis pupae are in diapause throughout a long winter period, and delayed emergence of the late population appears to be caused by prolonged diapause regulated by a developmental temperature threshold. The study has added information on the biology of turnip fly populations, a prerequisite for improved pest control.     

Timing of diapause termination in relation to variation in winter climate

In temperate insects, winters are typically endured by entering diapause, which comprises a deep resting stage. Correct timing of diapause termination is vital for synchronization of emergence with conspecifics and for mobilizing resources when conditions for growth and reproduction become favourable. Although critical to survival, the intrinsic and extrinsic drivers of diapause termination timing are poorly understood. In the present study, we investigate diapause development under a range of durations (10–24 weeks) spent at different temperatures (?2 to 10 °C) in the pupal diapausing butterfly Pieris napi Linnaeus (Lepidoptera:Pieridae). We determine: (i) the maximum cold temperature for diapause development; (ii) if pupae in diapause count cold days or cold sums; and (iii) whether diapause termination is distinct or gradual. The results indicate large and idiosyncratic effects of high and low nonlethal temperatures on diapause development in P. napi. Although all temperatures tested lead to diapause termination, a thermal optimum between 2 and 4 °C is observed. Lower temperatures lead to decreased eclosion propensity, whereas higher temperatures slow down development and increase emergence desynchronization. These data suggest that, rather than a simple cold‐summing process with a distinct diapause termination point, there are trade‐offs between time and temperature at the low and high end of the thermal range, resulting in a nonlinear thermal landscape showing a ridge of increasing eclosion propensity at moderate temperatures. The present study suggests that the effects of temperature on diapause development should be included in projections on post‐winter phenology models of insects, including pest species.     

Rapid and repeatable shifts in life‐history timing of Rhagoletis pomonella (Diptera: Tephritidae) following colonization of novel host plants in the Pacific Northwestern United States

Host shifts of phytophagous insect specialists to novel plants can result in divergent ecological adaptation, generating reproductive isolation and potentially new species. Rhagoletis pomonella fruit flies in eastern North America underwent a host shift ~160 ya from native downy hawthorn (Crataegus mollis) to introduced, domesticated apple (Malus domestica). Divergent selection on diapause phenology related to the earlier fruiting time of apples versus downy hawthorns resulted in partial allochronic reproductive isolation between the fly races. Here, we test for how rapid and repeatable shifts in life‐history timing are driving ecological divergence of R. pomonella in the Pacific Northwestern USA. The fly was introduced into the region via larval‐infested apples 40–65 ya and now attacks native black hawthorn (Crataegus douglasii) and introduced ornamental hawthorn (Crataegus monogyna), in addition to early‐ and late‐maturing apple varieties in the region. To investigate the life‐history timing hypothesis, we used a field‐based experiment to characterize the host‐associated eclosion and flight activity patterns of adults, and the feeding times of larvae at a field site in Vancouver, Washington. We also assessed the degree to which differences in host‐fruiting time generate allochronic isolation among apple‐, black hawthorn‐, and ornamental hawthorn‐associated fly populations. We conclude that host‐associated fly populations are temporally offset 24.4% to 92.6% in their seasonal distributions. Our results imply that R. pomonella possesses the capacity for rapid and repeatable shifts in diapause life history to match host‐fruiting phenology, which can generate ecologically based reproductive isolation, and potentially biodiversity in the process.     

Patterns of diapause frequency and emergence in swede midges of southern Ontario

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Over-wintering stage polymorphism of a bruchine beetle: geographical variation in optimal diapause strategy

Abstract.  1. Most temperate insects undergo diapause at a specific developmental stage to overcome severe winters. The bruchine beetle Bruchidius dorsalis in a warmer region in Japan, however, has diverse over-wintering stages – non-diapausing larvae, diapausing larvae, and diapausing adults, whereas in a cooler region, it over-winters only as the final instar larva or adult in diapause.
2. The geographical pattern of over-wintering stages in 12 populations was investigated over a wide range along the mainland of Japan. It revealed that a distinct geographical pattern of over-wintering stages exists in relation to temperature conditions. In regions with warmer climates, the proportion of non-diapausing larvae increased and B. dorsalis had a more complex over-wintering stage structure.
3. Life cycles were also compared between two areas of Japan by field experiments. In the cooler area, the first generation over-wintered in the diapausing larval or adult stage. Conversely, in the warmer area, diapause was induced later and some of the first-generation adults produced second-generation offspring before over-wintering.
4. Based on the geographical cline of climates and the differences in cold hardiness among stages, we can demonstrate that the over-wintering stage variation among and within populations results from an adaptive timing of diapause induction in each region, because the late larval or adult diapauses protect pupae or eggs – which unlike other stages are not cold hardy – from being produced late in the season.     

Quantitative trait locus mapping of resistance in apple to Cydia pomonella and Lyonetia clerkella and of two selected fruit traits

Apple, Malus×domestica, is the most important fruit grown within the temperate zonobiome. It is attacked by both fruit‐damaging and leaf‐damaging lepidopteran pest insects, which require regular control such as the carpophagous codling moth, Cydia pomonella, or frequent control such as the phyllophagous apple leaf miner, Lyonetia clerkella. As many environmentally friendly pest control tactics are only effective at low levels of infestation, host plant resistance is a promising future component of integrated pest management systems, but knowledge is still lacking on such genetically based approaches against lepidopteran pests. The aim of the study was to identify molecular markers linked to C. pomonella and L. clerkella resistance or susceptibility in commercial apple as well as markers linked to selected fruit traits. The number of C. pomonella‐infested fruits and the number of L. clerkella mines were quantified as measures of apple resistance or susceptibility to the studied moth species. Herbivore surveys on 160 apple genotypes, representing a segregating F₁ cross of the apple cultivars ‘Fiesta’ and ‘Discovery’, were carried out during two consecutive years and at two sites in Switzerland. Broad‐sense heritability was 29.9% (C. pomonella), 18.2% (L. clerkella), 21.9% (fruit number) and 16.6% (fruit diameter). A subsequent analysis identified a quantitative trait locus (QTL) associated to C. pomonella susceptibility on the Discovery linkage group 10. The closest marker to this QTL was the random amplified polymorphic marker Z19‐350. No significant QTL was identified for resistance to L. clerkella. A putative QTL associated to fruit number was identified on Fiesta linkage group 12. The presented QTL associated with C. pomonella susceptibility and the putative QTL linked to fruit number may facilitate marker‐assisted breeding of resistant apple cultivars with cropping traits desirable for optimal fruit production.     

Intra‐population genetic variation in diapause incidence of adult‐diapausing Tetranychus pueraricola (Acari: Tetranychidae)

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