Spermatogeny as a life-history index in parasitoid wasps

Reproduction is a major life-history trait but it has been studied mostly in relation to female reproductive effort. Recently, an ovigeny index using the proportion of oocytes ready to be oviposited at eclosion has been proposed for female insect parasitoids. Here, we propose a spermatogeny index for male parasitoid wasps. Prospermatogenic species have an index of 1, have all their spermatozoids mature at emergence and do not produce more later in life. At the other end of the spectrum, synspermatogenic species have no spermatozoids at emergence and produce them later in life. The level of spermatogeny should be linked to several other life-history parameters such as longevity, size, nutrition, distribution of mating opportunities and dispersion before and after mating. Data presented for some parasitoid species support the presence of variability in this male life-history parameter.     

Sex ratio response of the parasitoid wasp Muscidifurax raptor to other females

This study examines the sex ratio response of the parasitoid wasp Muscidifurax raptor to conspecific and confamilial females in relation to two groups of functional sex ratio models, local mate competition and host quality models. In some but not all experiments, M. raptor females produced a greater proportion of sons in the presence of a conspecific female than when alone, and this sex ratio effect carried over for a day after the females were isolated from each other M. raptor females also produced a greater proportion of sons in the presence of a female of the confamilial parasitoid Spalangia cameroni than when alone (although only on the second day of exposure to S. cameroni, not on the first). M. raptor's sex ratio increase in the presence of conspecifics is consistent with local mate competition models but not with host quality models because the presence of a conspecific female did not cause there to be more, and thus potentially smaller, offspring developing per host. In contrast, the presence of a S. cameroni female did cause there to be more offspring developing per host than when a M. raptor female was alone; thus M. raptor's sex ratio increase in the presence of S. cameroni may be explained by host quality models. An alternative explanation for the sex ratio increase in response to confamilials is that only a sex ratio response to conspecifics may be adaptive, due to local mate competition; but M. raptor females may be unable to distinguish between conspecific and S. cameroni females.     

高温胁迫对寄生蜂的影响及其适应机制

昆虫作为变温动物类群,极易遭受高温胁迫的影响,寄生蜂也不例外。寄生蜂是害虫生物防治领域中最常用的天敌昆虫,我国已将赤眼蜂广泛用于农林业鳞翅目害虫的防治。研究高温胁迫对寄生蜂的影响,可为全球气候变暖背景下农林害虫生物防治提供新的见解和对策。本文系统综述了高温胁迫对寄生蜂生命活动的影响以及寄生蜂响应高温胁迫的机制,并就温度升高对寄生蜂规模化繁育和应用可能出现的问题和挑战进行了展望,以期为更好地发挥寄生蜂在害虫生物防治中作用提供理论参考。     

COI DNA barcoding for molecular identification of the genus Aphidius (Braconidae: Aphidiinae) in South Korea,with diagnoses of two newly recorded species

The genus Aphidius is one of the most commercially important species in greenhouse for controlling pests, especially aphids. Although 13 Aphidius species had been recorded in South Korea, it was not carefully surveyed yet. In this study, we surveyed and collected 8 Aphidius spp. including two unrecorded species. Using the DNA barcode region of mitochondrial cytochrome c oxidase subunit I (COI), sequences were newly generated for 16 molecular operational taxonomic units (MOTUs) of 8 species, except for 15 duplicates, in addition of 58 MOTUs of 18 species referred from GenBank for comparison and identification, which were reconstructed in a barcode phylogeny with Neighbor-Joining method. Then, to define each species, both Automatic Barcode Gap Discovery (ABGD) and Bayesian Poisson Tree Processes (bPTP) analyses were performed. As results, although most of Aphidius spp. could be identified by clustering into a clade, we recognized that three Aphdius species were placed in a clade, of which each species was not separated as an independent species. Our data suggested that those species demonstrating a species complex were able to be hypothesized as one species. In addition, diagnoses and illustrations of the two newly recorded species in South Korea were provided.     

Haemocyte changes in D. Melanogaster in response to long gland components of the parasitoid wasp Leptopilina boulardi: a Rho-GAP protein as an important factor

The hymenopteran wasp Leptopilina boulardi (Figitidae) is a larval solitary parasitoid of Drosophila larvae of the melanogaster sub-group. The factors used by parasitoid females to prevent encapsulation of their eggs by the host are localized in the female long gland and reservoir. We report here the physiological effects of these factors on host haemocytes using in vivo injection experiments. The total number of haemocytes, the number of plasmatocytes and the number of crystal cells were not modified by injection of long gland extracts. In contrast, long gland extracts either from virulent or avirulent strains had a significant effect on the lamellocyte number. Compared to the Ringer control, the avirulent long gland products induced an increase of the lamellocyte number while virulent extracts induced a drastic decrease together with an alteration of the morphology of these cells. Interestingly, changes in the lamellocyte morphology were also observed following injection of the P4 protein, a major component of L. boulardi female long glands that displays a strong immune suppressive effect on Drosophila larvae. The implication of the P4 protein in suppressing the host cellular immunity is discussed in correlation with its predicted molecular function as a Rho-GAP protein.     

Temporal and spatial expression of the period gene in the reproductive system of the codling moth

The authors examined patterns of spatial and temporal expression of Drosophila per gene homologue in the codling moth, Cydia pomonella. Since sperm release in moths is regulated in a circadian manner by an autonomous clock that is independent from the brain, the authors investigated per expression in male reproductive system along with its expression in moth heads. per mRNA is rhythmically expressed with the same phase and amplitude in both tissues under light-dark (LD) conditions. The levels of per mRNA are low during the day, start to increase before lights-off, reach the peak in dark, and decrease after lights-on. In constant darkness (DD), cycling of per mRNA continued in heads with severely blunted amplitude. No cycling of per mRNA was detected in testis in DD. In situ hybridization and immunocytochemistry revealed distinct spatial patterns of per expression in the moth reproductive system. There is no expression of per in cells forming the wall of testes or in sperm bundles. However, per mRNA and protein are rhythmically expressed in the epithelial cells forming the wall of the upper vas deferens (UVD) and in the cells of the terminal epithelium, which are involved in the circadian gating of sperm release. Increase in per mRNA in the UVD coincides with sperm accumulation in this part of the insect reproductive system.     

Genetic inferences about the population dynamics of codling moth females at a local scale

Estimation of demographic parameters is important for understanding the functioning of natural populations and the underlying ecological and evolutionary processes that may impact their dynamics. Here, we used sibship assignment methods to shed light on the local dynamics of codling moth females in eight orchards in a 90-ha domain near Valence, France. Based on full-sib inference among 1,063 genotyped moths, we estimated (1) the effective number of females that had offspring, (2) their fertility and (3) the distribution of their oviposition sites within and among orchards. The average number of females in all the orchards increased between the first (~130) and the second (~235) annual generations. The average fertilities of the females were similar at each generation according to the host plant considered (apple, pear, or walnut), but differed between commercial (~10) and non-treated (~25) apple orchards. Females mainly clustered their eggs on contiguous trees along orchard borders, but they also occasionally dispersed their eggs among different orchards independently of the cultivated host plants or the inter-orchard distances (up to 698 m) during the second annual generation. The mean distance between two oviposition sites was 30 m. Sibship estimates of both the effective number of females and the inter-orchard migration rates (~5%) were in agreement with the observed genetic differentiation among the eight orchards (0.006 < F _st  < 0.013). These results confirm and extend previous field and laboratory observations in Cydia pomonella, and they demonstrate that sibship assignments based on genetic data are an interesting alternative to mark-release-recapture methods for inferring insect population dynamics.     

Biogenesis, structure, and immune-suppressive effects of virus-like particles of a Drosophila parasitoid, Leptopilina victoriae

Drosophila melanogaster larvae are attacked by virulent strains of parasitoid wasps. Females of Leptopilina heterotoma produce virus-like particles (VLPs) that efficiently destroy lamellocytes, a major larval immune effector cell type. We report here that L. victoriae, a closely related wasp species, also produces VLPs that trigger immune suppression responses in fly hosts. We compare the ability of immune suppression of the two parasitoids using a mutant host strain hopscotch^{Tumorous-lethal} (hop^Tum-l). hop^Tum-l larvae have two defects of hematopoietic origin: overproliferation of hemocytes and constitutive encapsulation of self-tissue by lamellocytes. The encapsulation phenotype is suppressed weakly by L. victoriae and strongly by L. heterotoma. In vitro studies on hop^Tum-l lamellocytes show that VLP-containing fluid from either wasp species induces lamellocyte lysis, but with different kinetics.Previously undocumented precursors of L. victoriae VLPs are synthesized in the long gland and are first visible within canals connecting secretory cells to the long gland lumen. VLP assembly occurs in the lumen. VLPs show multiple electron-dense projections surrounding a central core. Maturing particles appear segmented, singly or in arrays, embedded in the reservoir matrix. In sections, mature particles are pentagonal or hexagonal; the polygon vertices extending into spikes. Our results suggest that L. victoriae is likely to promote immune suppression by an active mechanism that is mediated by VLPs, similar to that used by L. heterotoma.     

新疆不同寄主植物苹果蠹蛾成虫动态调查

【背景】苹果蠹蛾是世界性重大检疫害虫,新疆是我国最早发现该虫的地区。监测苹果蠹蛾的种群动态能提高控制效率并阻止其在非疫区蔓延。【方法】利用性诱剂于2008~2009年期间监测苹果蠹蛾成虫在苹果园、梨园、李子园、桃园等生境下种群动态。【结果】苹果蠹蛾在北疆有2个明显高峰期,越冬代成虫高峰期为5月中旬至6月上旬,第1代成虫高峰期为7月中旬至8月中旬;在南疆有3个明显高峰期,分别为6月底至7月上旬、7月下旬至8月上旬和8月底至9月中旬。南疆和北疆的种群数量差异不是很大,单个性诱剂高峰期的种群数量小于10头,平均0.5~2.0头;同一区域苹果蠹蛾在不同作物之间存在一定的差异,但集中在苹果园和桃园。【结论与意义】新疆北疆苹果蠹蛾成虫一年一般发生2代,南疆3代。苹果蠹蛾以苹果园为主要发生区域,同一区域的苹果蠹蛾在相邻果园的苹果、葡萄、梨树上的种群动态具有密切相关性。苹果蠹蛾的种群动态为准确判定防治关键期提供依据,其在不同作物之间的相关性,为区域重点监测对象的判定提供基础依据。     

Below-ground host location by <Emphasis Type="Italic">Campsomeriella annulata</Emphasis> (Hymenoptera: Scoliidae), a parasitoid of scarabaeid grubs

Scoliid wasps are ectoparasitoids that attack soil-dwelling scarabaeid larvae, and little is known about their host-searching behavior. In this study we investigated the cues used in host location by Campsomeriella annulata (Fabricius) and examined whether or not these wasps can detect hosts in the soil from the surface. In a dual-choice test with a Y-tube, female wasps were attracted to sand with host odor, sand with host feces, and sand used for rearing the host, the larvae of Anomala rufocuprea Motschulsky (Coleoptera: Scarabaeidae). In a dual-choice test for cues presented at a distance, the wasps did not discriminate between the Y-tube arms with and without cues. In an experimental arena in which host products and a host grub were buried 0.5 cm below the surface the wasps did not respond to the cues from the surface in terms of the burrowing frequencies and antennal tapping rates. Our results indicate that C. annulata searches for the host grubs by using kairomones, residual cuticular substances, or feces deposited in the soil as the grubs move through it, and that wasps cannot perceive the host in the soil from the surface. We discuss how scoliid wasps search for soil-dwelling hosts using cues that are reliable but not highly detectable.     

Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth

Abstract. Mated female Brachymeria intermedia (Hymenoptera: Chalcididae) deprived, since emergence, from pupae of their host Lymantria dispar (Lepidoptera: Lymantriidae), accumulated eggs but had a very low rate of hostacceptance. Parasitoids that were host-deprived after encountering pupae early in life also accumulated eggs, but maintained a high acceptance rate. Thus early exposure to hosts promoted active reproductive behaviour. Total egg production depended on the total number of pupae encountered, indicating that B.intermedia adjust their egg production to host availability. Hence, in B. intermedia both the physiological state of the parasitoid (age and egg load) and the informational state (in this case host-availability and experience) interact to shape oviposition behaviour.     

Control of diapause in codling moth larvae

Diapause in a New Zealand strain of codling moth (Cydia pomonella Linnaeus [Lepidoptera: Olethreutidae]) was induced in larvae by photoperiods of 15 h or less. Once diapause had been initiated, it could not be terminated by any combination of conditions tested for at least 20 days after cocooning. In diapausing larvae a low rate of pupation occurred at 25 °C under a long day (18 h) photoperiod. A high rate of pupation was achieved under a long day regime when larvae were decocooned, and provided with apple as nourishment. Diapause could be terminated predictably in 94–100% of larvae by 1) conditioning at 15 °C and constant darkness for periods of 40–100 days, then 2) chilling at 2±2 °C and constant darkness for 20–50 days followed by 3) any post-chill condition periods at 25 °C, 18 h photoperiod. Complete diapause termination was achieved when 100 days conditioning was followed by 30 days or 50 days post-chill period. Under these conditions, 76% termination occurred in the post-chill period after 10 days, and 93% after 25 days.To terminate diapause in codling moth larvae, we recommend that a 100 days conditioning followed by 30 days chilling and 50 days post chilling periods be used.     

Structure,organization, and response of a species-rich parasitoid community to host leafminer population dynamics

The parasitoid community dynamics of an agromyzid honeysuckle leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera) were studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The parasitoid fauna composed three koinobionts (all larval-pupal solitary parasitoids) and 22 idiodiont species (11 larval solitary, nine pupal solitary and one pupal gregarious). The parasitoid community was dominated by early-attacking oligophagous braconid koinobionts at early periods, but was gradually displaced by late-attacking polyphagous eulophid idiobionts. Accordingly, the diversity index of the parasitoid community peaked at an intermediate point in the intra-generational succession. The succeeding attack-in-waves by the late-attacking idiobionts greatly reduced not only the survival rates of early-attacking parasitoid larvae but also the survival rates of hosts. The density-dependence observed in the host pupal mortality was thought to result from density-dependent host-switching by a keystone polyphagous pupal idiobiont parasitoid, Chrysocharis pubens, whereas high host pupal mortality was potentially attained by an early-attacking koinobiont braconid. Supposed aggregation of polyphagous parasitoids at high host density resulted in intense within-host competition and in an increase of host-feeding attack, both of which contributed to low emergence rates of parasitoids at high host densities. Parasitoid emergence rates were also reduced at low host densities, probably by inter- and intra-specific hyperparasitism among oligophagous parasitoids for limited hosts. The regulation effects of the species-rich parasitoid community upon the host population dynamics are thought to derive from succeeding attack-in-waves by polyphagous late-attacking idiobionts, especially by the keystone species.     

Commensal and mutualistic relationships of reoviruses with their parasitoid wasp hosts

During evolution, certain endoparasitoid wasps have developed mechanisms to suppress the defence systems of their hosts. For this purpose, these species, all of which belong to the families Ichneumonidae and Braconidae, inject various kinds of virus-like particles. The most studied of these particles are classified as polydnaviruses (family Polydnaviridae) which are symbiotic viruses. Over the past decade, it has also been shown that several wasp species harbour reoviruses (family Reoviridae), and that two of these suppress host defence, allowing the development of the parasitoid eggs. In this paper, we summarize the key features of these viruses and their relationships with their wasp hosts. Five reoviruses are known that appear to be non-pathogenic for the wasps. Three of these, McRVLP, HeRV, OpRVLP, use their wasp hosts as vectors, and do not appear to be involved in host defence suppression. The fourth, DpRV-1, is a commensal reovirus detected in most field populations of the wasp, Diadromus pulchellus. This reovirus is always found associated with an ascovirus, DpAV-4a, which is indispensable for host immune suppression. Although DpRV-1 has not been shown to directly increase D. pulchellus parasitic success, it may contribute to this success by retarding DpAV-4a replication in the wasp. The fifth reovirus, DpRV-2, occurs in a specific population of D. pulchellus in which DpRV-1 and DpAV-4 are absent. This virus has a mutualistic relationship with its wasp host, as its injection by females during oviposition is essential for host immunosuppression. Interestingly, these viruses belong to several different reovirus genera.     

Self and conspecific superparasitism by the solitary parasitoid Antrocephalus pandens

Abstract.

• 1 The fitness consequences of superparasitism for a solitary parasitoid depend on whether the host was first parasitized by itself (‘self-superparasitism’) or a different individual (‘conspecific superparasitism’). Self superparasitism is usually expected to be avoided.
• 2 A.pandens females showed no difference in their probability of superparasitism between self-parasitized and conspecifically-parasitized hosts. The probability of superparasitism decreased as time from the laying of the first egg in a host increased, from about 0.29–0.46 at a time interval of 1 h to 0.10–0.14 at 72 h.
• 3 The egg distribution of wasps foraging alone on a patch showed significant avoidance of superparasitism, but that of wasps foraging in the presence of conspecifics was not significantly different from a random distribution. This suggests that wasps switch from avoidance of superparasitism when alone to acceptance of all hosts when in a group.
• 4 When wasps foraged in a group, the hosts had many more ovipositor puncture marks than when wasps foraged singly. This suggests that either hosts were attacked several times per encounter, or that the wasps' encounter rate with hosts was much higher when in a group. If the latter is true, it is possible that, although the egg distribution suggested a higher rate of superparasitism when wasps foraged in a group, the ratio of acceptances to contacts of parasitized hosts may in fact have been lower.

     

Dispersal behavior of the parasitic wasp Cotesia urabae (Hymenoptera: Braconidae): A recently introduced biocontrol agent for the control of Uraba lugens (Lepidoptera: Nolidae) in New Zealand

Estimates of the dispersal range of a recently introduced biocontrol agent in its new environment are vital to understanding its relative searching capacity, and to foresee the maximum area that could be covered in a release event. In New Zealand, the solitary endoparasitoid Cotesia urabae Austin and Allen (Hymenoptera: Braconidae) was first released in January 2011 as a biological control agent for the gum leaf skeletoniser, Uraba lugens Walker (Lepidoptera: Nolidae). The objective of this study was to utilize an experimental approach to quantify the dispersal behavior of one generation of C. urabae. In our experiment, which used sentinel larvae as target hosts, parasitoids dispersed up to 20 m away from the release point but parasitism was highest within 5 m of the release site. A high level of parasitism was observed at the release tree (87.6%) which suggests that most of the females released may have stayed there. According to the dispersal model developed from the data collected, Cotesia would be able to disperse up to 53 m in one release event. In addition, significant differences were found between the different directions tested for dispersal, showing a clear downwind effect on dispersal suggesting that wind has a direct effect on the dispersal behavior of C. urabae in the field.     

Mortality factors affecting immature stages of codling moth,Cydia pomonella (Lepidoptera: Tortricidae), and the impact of parasitoid complex

Codling moth, Cydia pomonella Linnaeus (Lepidoptera: Tortricidae), is a serious pest of apples worldwide. This study aimed to evaluate the mortality rate of codling moth eggs, larvae and pupae in the field in commercial and neglected apple and walnut orchards over two years, and to investigate the biodiversity and intensity of parasitoids associated with codling moth in the orchards. Five patches of wax paper containing 1-day-old codling moth eggs were placed in a neglected orchard in order to evaluate parasitism rates. Corrugated cardboard bands were placed around the trunk of 15 trees during late spring and the beginning of summer through to fruiting season to capture and measure parasitism of codling moth larvae. 5285 larvae in total were collected during this study. Mortality rate (egg?+?larvae?+?pupae) varied between the commercial and neglected orchards, reaching a maximum of (42.89% and 66.67%) in neglected apple orchards and (61.03% and 74.76%) in the neglected walnut orchard in 2003 and 2004, respectively. Trichogramma cacoeciae (Hymenoptera: Tichogrammatidae) was the only egg parasitoid recorded. Eight hymenopteran larval and pupal parasitoids belonging to several subfamilies were recorded: Cheloninae, Agathidinae, Cremastinae, Haltichellinae, Chalcidinae, Anomalinae, and Pteromalinae and one dipteran belonging to Tachininae. In conclusion, mortality factors, mainly by parasitoids, are contributing to a general reduction in codling moth larvae populations particularly in neglected orchards. The hymenopteran Ascogaster quadridentata and the dipteran Neoplectops pomonellae can contribute to biological control programmes against codling moth in the coastal region and other regions.     

Production and storage of diapausing codling moth larvae

The rearing and production of diapausing codling moth (Cydia pomonella (L.) [Lepidoptera: Olethreutidae]) larvae for use in research programmes is outlined. Diapause was induced under conditions of complete darkness or under photoperiods of up to 15 h. Diapausing larvae were stored for over 2 years in the laboratory but greatest survival occurred when the storage time was one year or less. The recommended conditions for the induction of diapause are a 12 h photoperiod, 25°C and 55–65% r.h.

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Résumé Cet article porte sur les aspects principaux de l'élevage et de la production de chenilles diapaussantes de Cydia pomonella L. (Lep. Olethreutidae) pour des travaux de recherche. La diapause a été induite sous obscurité totale ou photophases inférieures à 15 heures. Des chenilles diapausantes ont été conservées en laboratoire jusqu'à deux ans, mais la survie pendant la première année était supérieure. Les conditions recommandées pour l'induction de la diapause sont une photophase de 12 heures, 25°C et 55–65 h.r.

     

Attraction of codling moth males to apple volatiles

The attraction of the codling moth, Cydia pomonella, to apple volatile compounds known to elicit an antennal response was tested both in the field and in a wind tunnel. In the field, (E)‐β‐farnesene captured male moths. The addition of other apple volatiles, including (E,E)‐α‐farnesene, linalool, or (E,E)‐farnesol to (E)‐β‐farnesene did not significantly augment trap catch. Few females were caught in traps which also caught male moths, but female captures were not significantly different from blank traps. In the wind tunnel, males were attracted to (E,E)‐farnesol, but not to (E)‐β‐farnesene. The addition of (E,E)‐α‐farnesene to (E)‐β‐farnesene had a synergistic effect on male attraction. The male behavioural sequence elicited by plant volatiles, including upwind flight behaviour, was indistinguishable from the behaviour elicited by sex pheromone.     

Effects of photophase entrainment on mating of the codling moth

Only 25.6% and 15.4% of adult male laboratory-reared diet-fed codling moths, Laspeyresia pomonella (L.), of a Polish strain mated in a field cage when they were reared from egg to adult with regimens of 1800–2400 or 2200–0400 hr darkness, respectively. However, 38.5%, 29.5%, and 29.4% of similar males mated (apple-reared from egg through larva in a 2200–0400 hr dark regime) after shifting the dark regimen in the late 5th instar to 1800–0400, 2200–0400 hr, or continuous dark, respectively.

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Résumé Une souche polonaise de Laspeyresia pomonella est élevée en laboratoire sur aliment artificiel, et soumise, de l'uf à l'adulte, à deux types de régime 1800–2400 h ou 2200–0400 h d'obscurité. Les adultes mâles issus de ces élevages et testés pour l'aptitude à l'accouplement dans des cages en nature, ne s'accouplèrent respectivement que dans la proportion de 25,6% et 15,4%. Toutefois on observe 38,5%, 29,5% et 29,4% d'accouplement avec des mâles comparables provenant d'élevages nourris sur pommes, et selon un régime de 2200–0400 h d'obscurité remplacé au cours du 5ème et dernier stade larvaire par 1800–0400, 2200–0400 h ou par l'obscurité continue.

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Financial support for this investigation was provided by the USDA under PL-480 Project PL-ARS-15 and the International Atomic Energy Agency Contract 13072. Mention of a proprietary product in this paper does not constitute an endorsement of this product by the USDA.