Field resistance of codling moth against Cydia pomonella granulovirus (CpGV) is autosomal and incompletely dominant inherited

The current appearance of local codling moth populations with resistance to Cydia pomonella granulovirus (CpGV) is an impediment to continuous CpGV application. Therefore, crossing experiments have been performed in order to gain information about the inheritance of resistance. Evidence is presented that the observed field resistance is stably inherited even under non-selective conditions in the laboratory. Offspring of reciprocal F(1) crosses between a susceptible ('S') and a resistant ('R') strain and backcrosses between F(1) and S were bioassayed at different CpGV concentrations. The resistant strain showed 100 times lower susceptibility in 7-day bioassays. The responses of the reciprocal crosses (male S x female R and female S x male R) did not differ significantly, indicating that resistance is autosomally inherited. The median lethal concentration for the F(1) progeny was intermediate between those of its parental strains. Mortality data obtained from the backcrosses suggested that inheritance of resistance is due to a non-additive, polygenic trait.     

Overcoming the resistance of codling moth against conventional Cydia pomonella granulovirus (CpGV-M) by a new isolate CpGV-I12

Recently, codling moth (CM, Cydia pomonella L.) populations with a significantly reduced susceptibility to C. pomonella granulovirus (CpGV) products have been observed in Germany. A novel CpGV isolate, designated CpGV-I12, is able to overcome the CpGV resistance. CpGV-I12 originated from Iran and showed superior efficacy in laboratory bioassays against a resistant CM strain (CpR), which has a 100-fold reduced susceptibility to commercially used isolate CpGV-M. Determination of the median lethal concentration (LC₅₀) indicated that CpGV-I12 is nearly as efficient in resistant CpR as CpGV-M in a susceptible CM strain (CpS). Beyond, CpGV-I12 caused superior mortality in CpS. Infection experiments showed that the resistance breaking effect can be observed in all instars of CpR. CpGV-I12 is a promising alternative control agent of CM in orchards where conventional CpGV products fail. In addition, we demonstrate in bioassays with recombinant expressed Cry1Ab that cross-resistance to CpGV and Bacillus thuringiensis products is not likely.     

Sex linkage of CpGV resistance in a heterogeneous field strain of the codling moth Cydia pomonella (L.)

The occurrence of codling moth populations in European apple orchards that were not controlled by Cydia pomonella granulovirus (CpGV) is the first reported case of field resistance against a baculovirus control agent. A monogenic dominant sex-linked mode of inheritance was previously demonstrated in single-pair crosses between a homogeneous resistant (CpRR1) and a susceptible (CpS) laboratory strain of codling moth. However, resistant field populations (CpR) are more heterogeneous in their levels of resistance, and the possibility that they could harbor different resistance genes to CpRR1 had not been directly addressed. Here we report single pair crossing experiments using a resistant codling moth strain collected from an apple orchard in the southwest of Germany. Single-pair crosses within the field strain revealed a genetic basis to the heterogeneity of CpR concerning CpGV resistance. Hybrid crosses to a susceptible laboratory strain and backcrosses of the F1 generation to the resistant CpR strain confirmed that the homogeneous CpRR1 and the heterogeneous field strain CpR share the same mode of inheritance. Thus the variable levels of CpGV resistance in field populations is likely due to frequency differences of the same resistance-conferring gene, rather than different genes, which will facilitate future efforts to monitor and manage resistance.     

Incidence of apple fruit and leaf surface metabolites on Cydia pomonella oviposition

Water soluble metabolites identified from surfaces of apple tree fruit and leaf stimulate oviposition in Cydia pomonella L. The effects of two artificial blends of primary metabolites representing fruit and leaf surfaces, respectively, and of components within the blends were examined on egg-laying after two time periods: 3 min and 25 min of darkness. An artificial mixture of six metabolites, viz., three sugar alcohols (sorbitol, quebrachitol, and myo-inositol) and three sugars (glucose, fructose, and sucrose) did stimulate egg-laying. Fructose, sorbitol, and myo-inositol are important components of the stimulatory blend. Contact durations may induce variations in egg-laying responses. After 3 min of darkness, there were no differences in numbers of females laying eggs nor in the numbers of eggs laid on cloths treated with the complete blends and the controls. There were, however, clear effects of groups of compounds and of individual compounds. Reduced blends without sugars and sugar alcohols were in many cases significantly less stimulatory than the complete blends and the controls. After 25 min of darkness, the proportions of females laying eggs as well as the numbers of eggs were higher after treatment with the complete blends than on the controls. The proportions of females laying eggs on cloths treated with the reduced blends were rather similar to the controls, whereas there were still significant effects on the numbers of eggs laid after treatment with reduced blends derived from fruits but not from leaves.     

Susceptibility of Cydia pomonella to Bacillus thuringiensis under laboratory and field conditions

The delta-endotoxin of 12 strains in 10 subspecies of Bacillus thuringiensis Berliner was highly active against Cydia pomonella (L.) when assayed under laboratory conditions on artificial diet. These results could not be confirmed in the field.The disappointing results obtained under field conditions are due to the behaviour of the target insect. C. pomonella larvae do not ingest food during penetration of the fruit. The larva bites pieces of the epidermis and deposits them without ingestion on top of the entry hole.

---

Zusammenfassung Das delta-Endotoxin von B.t. war in Laborversuch auf Kunstmedium gegenüber den Larven des Apfelwicklers, C. pomonella, sehr aktiv. Die hohe Aktivität konnte aber unter Feldbedingungen nicht mehr bestätigt werden.Es wurde nachgewiesen, dass die unbefriedigenden Resultate von B. thuringiensis unter Feldbedingungen auf das Verhalten der Junglarven zurückzuführen sind: Die Larven nehmen während dem Eindringen in den Apfel keine Nahrung auf, sondern deponieren die herausgebissenen Epidermistücke über der Einbohrstelle.

     

苹果蠹蛾气味结合蛋白基因CpomOBP20的克隆及表达分析

[目的] 通过克隆苹果蠹蛾气味结合蛋白CpomOBP20基因cDNA序列,分析其序列特征和表达谱,旨在更好地了解OBP基因在苹果蠹蛾生命活动过程中的作用,为该害虫的绿色防控提供理论支撑。[方法] 采用RT-PCR法扩增苹果蠹蛾气味结合蛋白CpomOBP20基因cDNA序列,并使用生物信息学软件对其核苷酸和氨基酸序列进行分析;基于qPCR技术分析CpomOBP20基因在苹果蠹蛾4龄幼虫不同组织(头、血淋巴、表皮、脂肪体、中肠、马氏管和唾液腺)以及雌雄成虫不同末端组织(头、触角、下唇须、喙、足和翅)中的表达情况,利用分子对接研究了CpomOBP20与3种保幼激素的结合能力。[结果] 苹果蠹蛾气味结合蛋白CpomOBP20基因的开放阅读框长459 bp,共编码152个氨基酸,等电点为6.30,蛋白分子质量为16.264 ku,N末端具有20个氨基酸组成的信号肽序列,蛋白质序列中具有6个保守的半胱氨酸残基,属于Classical OBP。序列分析表明,CpomOBP20的氨基酸序列与小菜蛾OBP (XP_011557123.1)的一致性最高,在亲缘关系上更加接近。qPCR结果表明,CpomOBP20基因在苹果蠹蛾4龄幼虫以及雌雄成虫不同组织中均有表达,其中在4龄幼虫的血淋巴中表达量最高,在雌雄成虫表达量最高的分别是翅和足,其次是头部。分子对接结果表明,CpomOBP20与3种保幼激素均具有较好的结合能力,可能参与保幼激素的结合与转运。[结论] 本研究明确了CpomOBP20的核苷酸和氨基酸序列的组成及编码蛋白的理化性质,并推测CpomOBP20的作用可能不仅局限于嗅觉识别,在非嗅觉器官中也可能起着重要的生理作用,为今后更深入地探究气味结合蛋白在苹果蠹蛾生命活动中的作用机理提供数据支撑。     

库尔勒香梨上苹果蠹蛾发生规律及其性信息素迷向防控效果

【目的】苹果蠹蛾是我国进境检疫性有害生物。库尔勒香梨是苹果蠹蛾在新疆果树种植区的主要寄主。研究苹果蠹蛾性信息素迷向防控技术在新疆库尔勒香梨产区的应用效果,可为该害虫的绿色防控提供参考。【方法】通过持续监测与调查,掌握苹果蠹蛾在库尔勒香梨上的发生规律和危害特点,并通过定期调查诱蛾量和蛀果率,比较微胶囊迷向剂、迷向丝、常规化学农药对苹果蠹蛾的防治效果。【结果】库尔勒香梨产区苹果蠹蛾一年发生2~3代,以老熟幼虫越冬,且死亡率较低。4月中旬越冬代羽化出土,5月开始为害;第1代成虫高峰期为7月上中旬,第2代成虫高峰期为8月下旬至9月中旬。微胶囊迷向剂、迷向丝、常规化学农药处理区每周的平均诱蛾量分别为2.85、2.98、4.95头·台~(-1),平均蛀果率分别为2.25%、0.8%、0.99%,与空白对照区的差异均达极显著水平。【结论】使用性信息迷向剂能够降低苹果蠹蛾第1代、第2代幼虫及成虫的数量。     

Field evaluation of commercial formulations of the codling moth granulovirus: persistence of activity and success of seasonal applications against natural infestations of codling moth in Pacific Northwest apple orchards

Inundative applications of the codling moth (CM), Cydia pomonella L., granulovirus (CpGV), which target neonate larvae before or during initial entry into fruit, offer potential for selective control of this key pest. In field tests on apple we compared the persistence and efficacy of single applications of three CpGV products approved for organic orchards in North America. In addition, the success of repeated (2–14) applications of one product (Cyd-X) as a principal control measure for CM in apple orchards was monitored following operational use by cooperating growers at four separate locations. In the first study, an early season application of all products at label rates remained highly effective for the first 24 h (averaging 94% larval mortality relative to controls) and moderately effective after 72 h (averaging 71% mortality) during dry sunny conditions. Significant activity remained up to 14 days, suggesting prolonged survival of the virus in UV-protected locations, such as the calyx of fruit. A second application later in the season was slightly less effective. Data obtained from commercial sites provide circumstantial evidence for the effectiveness of well-timed CpGV applications against CM outbreaks. In all cases where first generation larvae were targeted beginning at egg hatch (≈250 degree days) and treated areas monitored (0.3–1.6 ha plots), fruit damage during the second larval generation was reduced or eliminated. Based on the number of live larvae recovered throughout the season, mortality rates remained high (80.3–100% across sites). The cumulative number of moths caught in pheromone-baited traps was reduced (66–94%) in the second flight. Data from tree bands placed to catch diapause-destined larvae indicated overwintering generations remained low in treated sites (0.18 larvae/band).     

New method for testing solar sensitivity of commercial formulations of the granulovirus of codling moth (Cydia pomonella, Tortricidae: Lepidoptera)

A method for screening codling moth granulovirus (CpGV) formulation sensitivity to sunlight using specially prepared half apples and a solar simulator is described. The half apple preparation allows an even coverage of virus over the surface of the fruit that would not be possible using whole apples. Leaves and artificial medium were not usable for extended periods of exposure in the solar simulator due to excess drying. Fruit was sprayed with 10⁻³ and 10⁻⁵ dilutions of three commercial formulations of CpGV (Carpovirusine, Cyd-X, and Virosoft) and infested with codling moth neonates. Half of the sprayed fruit was exposed to 650 W/m² for 4 h in an Atlas Suntest CPS solar simulator resulting in an accumulated radiant energy of 9.36 × 10⁶ J/m² before they were infested with neonate codling moth larvae. Spraying non-irradiated fruit with the 10⁻³ dilution of Cyd-X and Virosoft resulted in nearly 100% mortality of neonate larvae. Irradiation reduced viral activity by 71-98% at the 10⁻³ dilution and by up to 32% at the 10⁻⁵ dilution relative to non-irradiated fruit. The procedures utilized enabled good preservation of the fruit throughout the incubation period and minimized invasion of the fruit by plant pathogens and saprophytic organisms. This laboratory method for screening candidate formulations and potential UV protectants could conserve time and resources by eliminating adjuvants with less potential in laboratory tests and field testing only the most promising candidates. It also enables year-round testing.     

Death of Cydia pomonella larvae and damage to apple fruit, after field application of codling moth granulosis virus

In field experiments, larvae of codling moth Cydia pomonella (L.) rarely acquired granulosis virus on hatching from the egg, but picked up most later, on the tree surface. Deposits of virus sprayed in 1.0% w/v skimmed milk did not affect neonate larval behaviour. Larvae died, usually in the first instar, after entering treated fruit, but they frequently entered via the calyx or near the base of the stalk or through cracks in the skin, where little feeding damage by first-and sometimes second-instar larvae was seen.

---

Résumé En verger, la pulvérisation d'oeufs de carpocapse avec du virus de la granulose en suspension dans l'eau (additionnée de lait écrémé dilué à 1%) n'a pas modifié la survie des chenilles avant pénétration dans le fruit; par contre la pulvérisation des arbres a provoqué une forte mortalité. Bien que des chenilles consommant des poils et la surface des feuilles aient été observées avant leur pénétration dans le fruit, ce qui aurait pu provoquer leur contamination par le virus, il semble que la contamination létale provienne des fruits seuls.La présence de produit n'a modifié ni le comportement larvaire, ni le taux de pénétration dans les fruits; la mortalité y a lieu ensuite, généralement au premier stade. Dans 74 à 78% des cas, les chenilles ont pénétré dans le fruit par le calice ou près de la base du pédoncule — aucun dégât provenant de larves du premier stade n'y était visible, de même que dans le calice pour les larves du deuxième stade. Par contre, toute pénétration par la surface du fruit était repérable dès le premier stade. Il est possible que la répartition des lieux de pénétration dans le fruit influe sur la létalité due au virus et explique les variations d'efficacité observées en verger. Un système de classification des dégâts, provoqués lors de la pénétration dans le fruit, de chenilles du premier au troisième stade est proposé pour évaluer l'efficacité des essais en verger.

     

Releases of Trichogramma platneri (Hymenoptera: Trichogrammatidae) in Apple Orchards under a Sterile Codling Moth Release Program

Based on the premise that augmented host numbers may help multiply and support parasitoid populations, the egg parasitoid Trichogramma platneri Nargarkatti was released in apple orchards which were participating in a sterile codling moth, Cydia pomonella (L.), release program. Nonviable eggs resulting from matings involving at least one sterile codling moth partner can be successfully parasitized by T. platneri. Grain moth-reared, as well as codling moth-reared, T. platneri were released either in the spring or in the spring and summer oviposition period(s) of wild codling moth. Sentinel codling moth eggs were hung weekly, for 3-day periods, from May until September to determine fluctuations in T. platneri populations both during and between releases. Low numbers of wild or nonviable codling moth eggs or other susceptible host eggs resulted in sufficient eggs to maintain low spring- or summer-introduced T. platneri populations. T. platneri reduced codling moth damage in trees in which the Trichogramma were released.     

The susceptibility of the pea moth, Cydia nigricana, to infection by the granulosis virus of the codling moth, Cydia pomonella

Laboratory studies demonstrated that neonate larvae of the pea moth, Cydia nigricana, are susceptible to infection with a granulosis virus (CpGV) isolated from the codling moth, Cydia pomonella. Comparative LC₅₀ values for C. nigricana and C. pomonella are 1.90 × 10⁵ and 1.54 × 10⁴ capsules/ml of diet, respectively. The virus extracted from CpGV-infected pea moth larvae is serologically related, and probably identical, to CpGV.     

Incidence and transmission of a granulovirus in a large codling moth [Cydia pomonella L. (Lepidoptera: Tortricidae)] rearing facility

Incidences of potential per os Cydia pomonella granulovirus (CpGV) transmission within a large codling moth colony were identified. CpGV was detected in the water which is used to wash egg sheets. When pre-neonates were extracted from eggs prior to emergence and tested for the presence of CpGV, 40% were found to carry amounts of CpGV detectable by a polymerase chain reaction (PCR) assay, suggesting possible transovarial transmission of the virus. Although symptoms typical of virus infection and larval death were found infrequently within communal rearing trays, the frequency with which CpGV DNA was detected by PCR assays increased from a mean of 31% of 10-day-old larvae to 94% of 25-day-old larvae. CpGV in codling moth cadavers remained virulent after being held at 60 degrees C for 3 days under conditions similar to the treatment of spent diet at the rearing facility before its disposal. PCR tests of surface samples taken from air filters and rearing rooms of the rearing facility were found to contain CpGV. Bioassays of surface samples from the diet trash bin and a filter through which outside air is passed before entering the rearing chambers resulted in significant codling moth neonate mortality. The virulence of CpGV in dust from the spent diet and the original inadvertent positioning of the diet trash bin directly below one of the air intake ducts are suggested as a possible additional source of CpGV contamination within the facility.     

Comparative activity of baculoviruses against the codling moth Cydia pomonella and three other tortricid pests of tree fruit

The granulovirus of Cydia pomonella (L.) (CpGV) offers potential for selective control of codling moth. Two major limitations of CpGV are its narrow host range and lack of persistence in the orchard agroecosystem. The nucleopolyhedroviruses of the alfalfa looper Autographa californica (Speyer) (AcMNPV) and those of the celery looper Anagrapha falcifera (Kirby) (AfMNPV) have broad host ranges. Comparative assays of CpGV, AcMNPV, and AfMNPV against codling moth neonate larvae revealed a 54-93-fold greater susceptibility of codling moth to the granulovirus than to the two nucleopolyhedroviruses based on the LC(50) values for each virus. The LC(50)s for CpGV, AfMNPV, and AcMNPV were 32.7 capsules/mm(2), 1.77 x 10(3) occlusion bodies (OBs)/mm(2), and 3.05 x 10(3)OBs/mm(2), respectively. The LT(50) determined for AfMNPV using an approximate LC(95) of the virus against neonate larvae was 3.6 days. Histological examination of tissues in moribund codling moth larvae that had been treated with AfMNPV revealed the presence of nonoccluded and unenveloped virus rods in midgut tissue. Neither OBs nor signs of infection were detected in other tissues. The activity of AfMNPV was also evaluated in three other tortricid apple pests (obliquebanded leafroller, Choristoneura rosaceana (Harris); Pandemis leafroller, Pandemis pyrusana Kearfott; and the oriental fruit moth, Grapholitha molesta (Busck)). Codling and Oriental fruit moths were significantly more susceptible to AfMNPV than were the two leafroller species.     

Effect of bisabolangelone on development ofCydia pomonella larvae and oviposition of the females

The effects of bisabolangelone on the development of codling moth larvae,Cydia pomonella L., and on ovipositional behaviour of the females were studied under laboratory conditions. Entry of the neonate larvae into apples and their development on a semi-synthetic medium were completely inhibited when the larvae were exposed respectively to 10 μg a.i./cm² and 20 μg a.i./ml of this compound. While concentrations of 1.25 to 5.0 μg a.i./ml bisabolangelone in the medium did not significantly affect larval development, exposure of the larvae to a higher rate (10 μg a.i./ml) resulted in 80% mortality during the first week. Nevertheless, the larvae which survived the treatments underwent further development until emergence of the adults. No significant changes in duration of larval or pupal periods were recorded. Oviposition of the females in plastic beakers, whose inner surfaces were partially painted with two concentrations of bisabolangelone (1.25 to 5.0 μg a.i./cm²), was significantly reduced and the eggs were mainly laid on the parts painted with the ethanol solvent alone. When the inner surface of the cups was completely treated (i.e. top, bottom, and side) with similar concentrations of bisabolangelone, a dramatic reduction in oviposition occurred and the eggs were mostly laid on the bottom of the beakers. While this compound did not significantly influence egg hatch, it inhibited the normal upward movement of the adults in the cups and migration of the newly hatched larvae through perforations of the lids.     

Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.     

Identification of further sex pheromone synergists in the codling moth, Cydia pomonella

A number of angiosperm nonhost volatiles (NHVs) and green leaf volatiles (GLVs) were tested alone and as supplements to the antiaggregation pheromone, verbenone, for their ability to disrupt attack by the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae), on lodgepole pine, Pinus contorta Dougl. ex Loud. var. latifolia Engel. Preliminary experiments led to a refined NHV blend [benzyl alcohol, guaiacol, benzaldehyde, nonanal, salicylaldehyde, and conophthorin] and a refined GLV blend [(Z)-3-hexen-1-ol and (E)-2-hexen-1-ol]. In a 20-replicate experiment, NHV and GLV groups both singly, and verbenone alone, significantly reduced MPB mass attack on pheromone-baited trees and on trees within 5 m of the pheromone-baited trees. Both blends in combination with verbenone reduced the number of mass attacked, baited trees to three out of 20 compared to 20 out of 20 of the baited controls. Each binary combination was also effective at reducing mass attack. In these experiments, all tested repellents were released from devices stapled to trees at the same point as the pheromone bait, suggesting that the repellency could have been to a point source, rather than to the whole tree. Therefore, in two further experiments bands of release devices were wrapped around the treated trees and the pheromone bait was removed from the treated trees. In one experiment, when the aggregation pheromone bait was suspended between pairs of trees treated with the NHV blend plus GLV blend plus verbenone, only three out of 25 treated pairs had mass attack on at least one member of the pair. In the other 60-replicate experiment, with no pheromone baits present, attack occurred on 13 untreated and 11 banded trees, all in the path of a large advancing infestation. However, the mean attack density on the banded trees was significantly reduced to a level below the 40 attacks m^–2 of bark surface required to kill a healthy lodgepole pine. As a result of these experiments, operational trials are recommended.     

松瘿小卷蛾及其寄生蜂对落叶松挥发物的嗅觉行为反应

为确定松瘿小卷蛾（Cydia zebeana）对寄主的定位习性,以及探讨对其的综合防治方法,通过Y型管嗅觉仪行为选择实验,分析松瘿小卷蛾及其两种寄生性天敌小卷蛾革腹茧蜂（Ascogaster olethreuti）和卷蛾长体茧蜂（Macrocentrus sp.）对寄主兴安落叶松（Larix gmelinii）的9种不同浓度挥发物的行为反应。结果表明,对雌蛾产生引诱作用的挥发物为3-蒈烯、水芹烯,产生驱避作用的挥发物为月桂烯、罗勒烯、莰烯;（R）-α-蒎烯表现为低浓度引诱高浓度驱避。对雄蛾产生引诱作用的挥发物为（S）-β-蒎烯,产生驱避作用的挥发物为月桂烯、3-蒈烯、（S）-α-蒎烯、莰烯。对小卷蛾革腹茧蜂有引诱作用的挥发物为（S）-β-蒎烯、（S）-α-蒎烯、叶醇,产生驱避作用的挥发物为罗勒烯、莰烯、水芹烯;对卷蛾长体茧蜂有引诱作用的挥发物为（S）-β-蒎烯、（S）-α-蒎烯、（R）-α-蒎烯、叶醇,产生驱避作用的挥发物为罗勒烯、水芹烯。说明在利用推拉策略防控松瘿小卷蛾时,月桂烯、叶醇和（S）-α-蒎烯可用于保护目标林,驱避害虫吸引天敌;水芹烯可以结合化学防治施用于饵木林,诱杀害虫而驱避寄生蜂。     

Termination of the facultative diapause in the codling moth, Laspeyresia pomonella (Lepidoptera, Tortricidae)

The abiotic factors regulating the termination of the facultative diapause of the mature larva of the codling moth, Laspeyresia pomonella (L.), are described. The termination of diapause under long-day conditions (LD) is influenced by: (i) the rearing temperature of the larvae during prediapause development, (ii) the duration of the preincubation period, i.e. the time for which the diapausing larvae remain under prediapause rearing conditions, (iii) the reactivating incubation, i.e. the period for which the larvae are chilled, and (iv) the complementary incubation (LD and, except in one experiment, 26°), i.e. the period after the chilling needed for the pupation of the insects. The complementary incubation is distinctly shorter if the prediapause development of the larvae takes place at rearing temperatures below 26°, i.e. 21° or 19°. The latter conditions led to 100% pupae and pupation could be accelerated by prolonging the preincubation period. In larvae reared at 26°, the prolongation of the preincubation period raised the rate of pupation to a maximum of only 56% and also caused higher mortality. On the other hand, diapause was terminated under short-day conditions if the temperature was raised to 26° after a rearing temperature of 19° and a preincubation period of 90 days.

---

Beendigung der fakultativen diapause des apfelwicklers, Laspeyresia Pomonella (Lepidoptera, Tortricidae)

Zusammenfassung Es wurde untersucht, welche abiotischen Faktoren die Beendigung der fakultativen Diapause ausgewachsener Apfelwicklerlarven (Laspeyresia pomonella) steuern. Die Beendigung der Diapause unter Langtagbedingungen (LT) wird beeinflusst durch: 1. die Zuchttemperatur der Larven während der Prädiapauseentwicklung; 2. die Dauer der Präinkubation, d.h. die Zeit, während der diapausierende Larven unter Prädiapause-Zuchtbedingungen bleiben; 3. die Reaktivierungs-inkubation, d.h. eine Periode von 70 d, während der die Larven auf 4° gekühlt werden und 4. die Komplementärinkubation (LT und, ausser in einem Experiment, 26°), d.h. die nach der Kühlung bzw. dem Wechsel in der Photoperiode benötigte Zeitdauer bis zur Verpuppung. Die Komplementärinkubation ist deutlich kürzer, wenn die Prädiapauseentwicklung der Larven bei Temperaturen unter 26° stattfindet. Bei 19° verpuppten sich 100% der Larven, wobei der Zeitpunkt der Verpuppung durch die Verlängerung der Präinkubationszeit beschleunigt wurde. Bei Larven, die bei 26° gezüchtet wurden, erreichte die Verpuppungsrate lediglich 56% bei einer relativ langen Präinkubationsdauer, wobei deren Verlängerung auch die Larvenmortalität erhöhte. Nach einer Zuchttemperatur von 19° und einer Präinkubationsdauer von 90 Tagen konnte die Diapause unter Kurztagbedingungen beendet werden, wenn die Temperatur auf 26° erhöht wurde.

     

Does aggregation behavior of codling moth larvae,Cydia pomonella,increase the risk of parasitism by Mastrus ridibundus?

Mastrus ridibundus is a specialist hymenopteran parasitoid that parasitizes last-instar larvae or prepupae of the codling moth, Cydia pomonella. Foraging females eavesdrop on an aggregation pheromone produced by cocooning larvae. We investigated whether larvae that cocoon in aggregation experience a greater rate of parasitism than larvae that cocoon in isolation. In wind tunnel experiments, 10 larvae in aggregations were more readily located by female M. ridibundus than 10 larvae well separated from each other. Similarly, aggregations of 30 larvae were more attractive to female M. ridibundus than those of 3 larvae. In cage experiments, larval cocooning in aggregation or isolation had no effect on the mean rate of parasitism and the mean number of eggs deposited per parasitized host. In Petri-dish experiments, the location of larvae within an aggregation significantly affected their rate of parasitism, with those in the center of an aggregation completely shielded from parasitism. Our data suggest that aggregation behavior by C. pomonella larvae does not appear to increase the rate of parasitism. The increased risk of aggregated larvae to be detected by M. ridibundus is likely offset by diluted parasitism risk and structural refugia effects that larvae in aggregation experience. As an egg-limited parasitoid, female M. ridibundus can parasitize on average only one larva in an aggregation, with the likelihood of parasitism for each larva being inversely proportional to the number of larvae in that aggregation.