No loss of genetic divergence in Torymus sinensis following its release for Dryocosmus kuriphilus control in Europe

1. Arrival, establishment, and further dispersal of non-native natural enemies are considered essential for a successful biological control programme, while among the factors that may determine the success of such a programme, genetic diversity of the introduced population plays an important role in the establishment of a non-native species.
2. The Chinese parasitoid wasp Torymus sinensis Kamijo (Hymenoptera: Torymidae) was initially released in Europe in Italy to control biologically the Asian chestnut gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), and reduce the damage induced on sweet chestnut (Castanea sativa Miller). In the following years, T. sinensis was then released in numerous other European countries as a biological control agent of D. kuriphilus. Its presence has also been reported beyond the countries of release due to rapid natural dispersal.
3. To assess the post-release genetic diversity of D. kuriphilus, we screened T. sinensis populations from six European countries and tested the possibility of these populations suffering from frequently observed genetic effects that could threaten its successful establishment in Europe.
4. Our results exhibit that T. sinensis populations have suffered neither from the Allee effect nor from genetic bottleneck after their release and establishment in Europe, something that increases the possibility to effectively control D. kuriphilus in Europe.

     

Rearing,release and settlement prospect in Italy of <Emphasis Type="Italic">Torymus sinensis</Emphasis>, the biological control agent of the chestnut gall wasp <Emphasis Type="Italic">Dryocosmus kuriphilus</Emphasis>

Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), is one of the most serious pests attacking chestnut trees. Recently it was incidentally introduced into Northwest Italy and it is now spreading throughout Europe. This pest was successfully controlled in Japan by introducing a parasitoid, Torymus sinensis Kamijo (Hymenoptera: Torymidae), from China’s mainland. Following this successful experience, the parasitoid was introduced into Italy from Japan. One year of preliminary studies led to a successful method of rearing imported galls with a synchronization between the parasitoid’s emergence and the presence of the target galls in the field. In two consecutive years, a total of 2,117 individuals were released in several sites covering most of the infested area. There are encouraging data about the settlement of the parasitoid and its synchrony with the host’s cycle: from about 64,000 host galls collected in the field over 200 T. sinensis were reared. The role of native parasitoids associated with the chestnut gall pest in its Italian distribution range is also discussed.     

Influence of the introduced parasitoid Torymus sinensis (Hymenoptera: Torymidae) on T. koreanus and T. beneficus as indigenous parasitoids of the chestnut gall wasp Dryocosmuskuriphilus (Hymenoptera: Cynipidae) on chestnut trees in Nagano Prefecture, Japan

We describe here the parasitoid wasps Torymus sinensis Kamijo and T. beneficus Yasumatsu & Kamijo (early-spring and late-spring strains), which are introduced and indigenous natural enemies of the chestnut gall wasp Dryocosmus kuriphilus Yasumatsu, an invasive chestnut pest in Japan. We recently discovered specimens of T. koreanus Kamijo, endemic in Korea, among Torymus parasitoids collected from D. kuriphilus galls in a Japanese chestnut orchard. In this study we compare the composition of Torymus parasitoids emerging from D. kuriphilus galls before and after the release of T. sinensis. Before the release of T. sinensis, early-spring and late-spring strains of T. beneficus predominated (58.3 and 20.8% of specimens collected). However, a few years after the release, both T. beneficus strains had been almost completely displaced by T. sinensis. In contrast to the rapid decrease in T. beneficus, T. koreanus did not decrease drastically before and even after the release of T. sinensis (approximately 10–20% of specimens collected). These results suggest that not a few T. koreanus were present in the Japanese chestnut orchard investigated at least several years after the release of T. sinensis, although both the T. beneficus strains were rapidly displaced by T. sinensis during this period.     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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<Emphasis Type="Italic">Torymus sinensis</Emphasis>: a viable management option for the biological control of <Emphasis Type="Italic">Dryocosmus kuriphilus</Emphasis> in Europe?

The chestnut gall wasp Dryocosmus kuriphilus is a global pest of chestnut (Castanea spp). Established as a pest in the mid-twentieth century in Japan, Korea and North America, this species was first reported in Europe in 2002. Following the successful release of a biological control agent Torymus sinensis in Japan, this parasitoid species has been released in Italy since 2005. Here we discuss the potential of T. sinensis as a viable management option for the biological control of D. kuriphilus in central Europe. We suggest that more consideration should be given to determining, (i) the conditions under which T. sinensis may attack alternative native gall wasp hosts and (ii) the likelihood of hybridization of this species with native Torymus. Both issues are central to predicting unassisted range expansion by released T. sinensis, and to assess the environmental risks associated with a more widespread release of this species in Europe.     

Environmental thermal levels affect the phenological relationships between the chestnut gall wasp and its parasitoids

Studies of thermal level‐related asynchrony in a host–parasitoid relationship are necessary to understand the effects of climate change on new host–parasitoid interactions. In the Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae) and its Chalcidoidea parasitoids, phenological synchrony is assumed to be weather‐dependent in a new area of expansion. To evaluate the effects of environmental thermal regimes on the host, a phenology model for different cynipid stages (larvae, pupae, adults, and adult emergence) and a host–parasitoid phenological estimator are developed in three chestnut fields during two successive growth seasons and subsequently validated in areas with chestnut fields at two different altitudes. Comparisons of the timings of the juvenile and adult stages with those of the parasitoid complex demonstrate that the shortest period of occurrence for cynipids within galls has negative effects on the host–parasitoid relationships at higher temperature levels, thereby increasing phenological asynchrony for some parasitoids species. Reducing the development time of pupae and adults decreases the likelihood of success for some parasitoid species at higher temperature levels. We also record the extension of the gall wasp development time (approximately 15 days) at higher altitudes (linked to a lower mean temperature of approximately 1.5 °C). These results highlight how parasitization on the new hosts is dependent on the host phenology and, in the present study, is limited by the short duration of the presence of the host in galls, which could explain the considerable differences in cynipid gall wasp parasitization recorded at different altimeters.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

Identification of Torymus sinensis and T. beneficus (Hymenoptera: Torymidae), introduced and indigenous parasitoids of the chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae), using the ribosomal ITS2 region

The parasitoid wasps Torymus sinensis and T. beneficus (Hymenoptera: Torymidae) are introduced and indigenous natural enemies, respectively, of the chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae), an invasive pest of chestnuts in Japan. T. beneficus has two emergence types in spring, here tentatively designated as the early-spring strain and late-spring strain. It is very difficult to distinguish these two Torymus species accurately according to their morphological and ecological characteristics. Although the sequences of internal transcribed spacer 2 of the nuclear ribosomal DNA (ITS2) of these parasitoids are very similar, we have developed a pair of primers for amplifying a part of ITS2 that distinguishes species and emergence type. By performing high-resolution electrophoresis of PCR products amplified by this specific primer pair, we have succeeded in accurately identifying T. beneficus (early-spring strain) and some T. sinensis parasitoids. However, this technique was discovered to be inapplicable to identify other T. sinensis and T. beneficus (late-spring strain) parasitoids with the same ITS2 genotype. In spite of this problem, the ITS2 marker appears to be more powerful than any other molecular markers so far reported, since it is applicable to detection of T. beneficus, T. sinensis, and their hybrids.     

Wolbachia prevalence patterns: horizontal transmission,recombination, and multiple infections in chestnut gall wasp-parasitoid communities

The Oriental chestnut gall wasp, Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), is a global invasive pest that causes serious damage to almost all chestnut species belonging to the Castanea genus (Fagaceae). Dryocosmus zhuili Liu et Zhu is a recently described sibling species of D. kuriphilus, which induces galls on Castanea henryi (Skan) Rehd. et Wils. There are many indigenous parasitoid species in China which play an important role in the natural regulation of their population dynamics. Wolbachia is a maternally inherited α-proteobacterium widely found in arthropods. This study screened for the presence of Wolbachia in the two chestnut gall wasps and in six parasitoid species from 12 populations, to investigate the prevalence patterns of Wolbachia in the chestnut gall wasp-parasitoid communities. We found that D. zhuili and four parasitoid species were infected with Wolbachia; among them, all individuals of the two populations of Megastigmus sp. had multiple Wolbachia infections. By using multilocus sequence types to characterize bacterial strains, three new sequence types were identified. The Wolbachia strains infecting D. zhuili (ST-507), Torymus sinensis Kamijo (ST-508), and Sycophila variegata (Curtis) (ST-508) belonged to supergroup A, whereas the Wolbachia strain infecting Megastigmus nipponicus Kamijo (ST-503) belonged to supergroup B. Our results also suggested that horizontal transmission of Wolbachia occurs between chestnut gall wasps and their parasitoids. Moreover, multiple Wolbachia infections of Megastigmus sp. may be due to gene recombination and horizontal transmission.     

Seasonal flight activity and genetic relatedness of Torymus species in Italy

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Rapid on-site identification of the biocontrol agent of the Asian chestnut gall wasp

In classical biocontrol programmes, a rapid and correct identification of the introduced antagonist is a key issue during both the release and establishment monitoring phases. It is often difficult to distinguish morphologically cryptic species or immature stages. An accurate diagnosis can now be provided by molecular diagnostic methods. Among the conventional and real-time PCR-based methods, loop-mediated isothermal amplification (LAMP) is a particularly suitable technique as it allows a rapid amplification of target DNA directly in the field. During the programme implemented in Italy against the Asian chestnut gall wasp (ACGW) Dryocosmus kuriphilus, we developed a real-time LAMP assay, combined with a simple DNA extraction method, for rapid in-field identification of larvae, pupae, and adults of the biocontrol agent, the parasitoid Torymus sinensis. Validation of the assay comprised adults as well as preimaginal stages of parasitoids obtained from ACGW galls collected from different localities. Results confirmed the effectiveness of the LAMP assay to rapidly and specifically identify the target parasitoid in the field. This assay will be a valuable tool for quick on-site checking of the parasitism rate.     

Success of classical biocontrol agent Torymus sinensis within its expanding range in Europe

Torymus sinensis Kamijo (Hymenoptera, Torymidae), a classical biocontrol agent of chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera, Cynipidae), was released in Croatia, Slovenia and Hungary in 2015. Following the introduction, the research of parasitism rates and population genetic indices on 40 different sites was performed to monitor and evaluate the success of establishment. The observed parasitism rates were unexpectedly high and negatively correlated with the distance from westernmost locality, while population genetic indices showed that populations of T. sinensis did not suffer from bottleneck‐induced founder effect phenomenon. Moreover, lack of genetic differentiation demonstrates that all populations share similar genetic structure, which could be shaped only by high levels of gene flow. We conclude that T. sinensis established viable and genetically diverse populations and successfully spread naturally from Italy across Slovenia to Croatia and Hungary.     

Non-random distribution among a guild of parasitoids: implications for community structure and host survival

Abstract 1. Immature stages of the gall midge, Asphondylia borrichiae, are attacked by four species of parasitoids, which vary in size and relative abundance within patches of the gall midge’s primary host plant, sea oxeye daisy (Borrichia frutescens). 2. In the current study, a bagging experiment found that the smallest wasp, Galeopsomyia haemon, was most abundant in galls exposed to natural enemies early in the experiment, when gall diameter is smallest, while the wasp with the longest ovipositor, Torymus umbilicatus, dominated the parasitoid community in galls that were not exposed until the 5th and 6th weeks when gall diameter is maximal. 3. Moreover, the mean number of parasitoids captured using large artificial galls were 70% and 150% higher compared with medium and small galls respectively, while stem height of artificial galls significantly affected parasitoid distribution. Galls that were level with the top of the sea oxeye canopy captured 60% more parasitoids compared with those below the canopy and 50% more than galls higher than the plant canopy. 4. These non‐random patterns were driven primarily by the differential distribution of the largest parasitoid, T. umbilicatus, which was found significantly more often than expected on large galls and the smallest parasitoid of the guild, G. haemon, which tended to be more common on stems level with the top of the plant canopy. 5. Large Asphondylia galls, especially those located near the top of the Borrichia canopy, were more likely to be discovered by searching parasitoids. Results using artificial galls were consistent with rates of parasitism of Asphondylia galls in native patches of sea oxeye daisy. Gall diameter was 19% greater and the rate of parasitism was reduced by almost 50% on short stems; as a result, gall abundance was 24% higher on short stems compared with ones located near the top of the plant canopy. 6. These results suggest that parasitoid community composition within galls is regulated by both interspecific differences in ovipositor length and preferences for specific gall size and/or stem length classes.     

Post-release evaluation of non-target effects of <Emphasis Type="Italic">Torymus sinensis</Emphasis>, the biological control agent of <Emphasis Type="Italic">Dryocosmus kuriphilus</Emphasis> in Italy

A post-release study was performed to assess the impact of Torymus sinensis (Hymenoptera: Torymidae), the biological control agent of Dryocosmus kuriphilus (Hymenoptera: Cynipidae), on native cynipid gall inducers in Italy. In total, 14,512 non-target galls were collected, corresponding to seven genera: Andricus, Aphelonyx, Biorhiza, Cynips, Diplolepis, Neuroterus, and Synophrus, and 8708 chalcid parasitoids were recorded. The Torymidae family accounted for about 30%, and Bootanomyia (=Megastigmus) dorsalis, Torymus affinis and T. flavipes were the most represented species. A total of 116 T. sinensis emerged from 15 different oak galls, mainly Andricus curvator and A. inflator. In controlled conditions, oviposition was recorded on A. cydoniae, A. grossulariae and A. lucidus, while no mating with native congeneric species occurred. This paper confirms the realised host-range expansion by T. sinensis. Even if it were extremely difficult to evaluate its magnitude, the impact appears minimal, and an occasional feeding with no changes in the distribution or abundance of non-target hosts is expected.     

Biology and monitoring of Dryocosmus kuriphilus on Castanea sativa in Southern Italy

• 1 The cynipid gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) is an invasive wasp in Europe and is one of the major pests of chestnuts worldwide. An investigation of this cynipid was started in Campania, Southern Italy, in 2008, with the aim of providing guidelines for its sustainable control.
• 2 The number of larvae/gall increased in the second year of colonization relative to the first one but thereafter decreased in the third year of colonization, whereas the volume of the galls after the initial increase in the second year of colonization became stable. Adult emergence generally peaked between the middle of June and July.
• 3 The only reliable method for assessing the presence of the pest at its earliest stage was through dissecting the buds because oviposition scars on the buds tended to disappear during the winter.
• 4 Yellow traps were shown to be an effective method for assessing the emergence of cynipid adults and could potentially be used to time chemical applications in those cases in which this control method is feasible.
• 5 No correlation was recorded between the relative durations of the final stages (pupae, pharate adults and emergence).
• 6 Possible control strategies for this invasive pest are discussed.

     

Plant neighbour identity and invasive pathogen infection affect associational resistance to an invasive gall wasp

Theory predicts that mixed forests are more resistant to native pests than pure forests (i.e. associational resistance) because of reduced host accessibility and increased top-down control by natural enemies. Yet, whether the same mechanisms also apply to invasive pests remains to be verified. We tested the hypothesis of associational resistance against the invasive Asian chestnut gall wasp (ACGW, Dryocosmus kuriphilus) by comparing ACGW infestation rates on chestnuts (Castanea sativa) in stands varying in species composition (chestnut alone or associated with oaks, pines or ashes). We investigated the effects of reduced chestnut density and frequency in mixed stands, as well as the effect of biotic interactions between ACGW, its parasitoids and the chestnut blight disease (caused by Cryphonectria parasitica). ACGW infestation rates were significantly lower in chestnut–oak and chestnut–ash mixtures than in pure chestnut stands and chestnut–pine mixtures. Infestation rate decreased with decreasing chestnut relative proportion. The composition of native parasitoid communities emerged from galls significantly differed between pure and mixed chestnut stands, but not the species richness or abundance of parasitoids. The abundance of the introduced parasitoid Torymus sinensis was not correlated with ACGW infestation rates and was independent of stand composition. Blight symptoms modified ACGW infestation rates with taller trees being preferred when they were asymptomatic but avoided when they presented blight disease damage. Our results suggest that conservation biological control based on tree species mixtures could contribute to reducing the damage of invasive forest pests.     

Invasion by the chestnut gall wasp in Italy causes significant yield loss in Castanea sativa nut production

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The parasitoid community of <Emphasis Type="Italic">Andricus quercuscalifornicus</Emphasis> and its association with gall size,phenology, and location

Plant galls are preyed upon by a diverse group of parasitoids and inquilines, which utilize the gall, often at the cost of the gall inducer. This community of insects has been poorly described for most cynipid-induced galls on oaks in North America, despite the diversity of these galls. This study describes the natural history of a common oak apple gall (Andricus quercuscalifornicus [Cynipidae]) and its parasitoid and inquiline community. We surveyed the abundance and phenology of members of the insect community emerging from 1234 oak apple galls collected in California’s Central Valley and found that composition of the insect community varied with galls of different size, phenology, and location. The gall maker, A. quercuscalifornicus, most often reached maturity in larger galls that developed later in the season. The parasitoid Torymus californicus [Torymidae] was associated with smaller galls, and galls that developed late in the summer. The most common parasitoid, Baryscapus gigas [Eulophidae], was more abundant in galls that developed late in the summer, though the percentage of galls attacked remained constant throughout the season. A lepidopteran inquiline of the gall (Cydia latiferreana [Tortricidae] and its hymenopteran parasitoid (Bassus nucicola [Braconidae]) were associated with galls that developed early in the summer. Parasitoids and inquilines, in general, had a longer emergence period and diapause within the gall than the gall-inducer. The association of different parasite species with galls of different size and phenology suggests that different parasite species utilize galls with slight differences in traits.     

Alien herbivores and native parasitoids: rapid developments and structure of the parasitoid and inquiline complex in an invading gall wasp Andricus quercuscalicis (Hymenoptera: Cynipidae)

Abstract.

• 1 Rapid and substantial changes have occurred in the parasitoid and inquiline community associated with the agamic galls of Andricus quercuscalicis since it invaded Britain in the late 1950s. The number of parasitoid and inquiline species has risen from one to thirteen over a 15-year period. Although the number of species has been relatively consistent over the last 8 years, the species composition has changed considerably and in a highly characteristic way during this period.
• 2 The parasitoid complex can be divided into two broadly distinct sets of parasitoid species; one set attacks only the gall former whereas the other set concentrates on the inquilines living in the wall of the gall.
• 3 The most dramatic change, however, is in the abundance of inquilines which were reported to be virtually absent in earlier studies on this community in Britain. Over a period of only 5 years, between 1988 and 1993, inquiline attack rose from less than 0.01 to an average of 0.26 inquilines per gall. The intensity of inquiline attack is geographically heterogenous, with high inquiline numbers restricted to south-east England. Because of the relatively high specificity of the parasitoids, high inquiline abundance is positively correlated with parasitoid species richness in knopper galls.
• 4 Parasitism rates, particularly on the gall former, were generally low (<10%). Over the last 5 years, however, seven parasitoid species have been consistently recorded and the mortality caused by these species has increased continuously. The species composition of the community associated with this alien gall wasp in Britain has quickly converged to the community known from its native range in continental Europe. Parasitoid species known to attack the galls of A.quercuscalisis on the continent have been recorded from it in Britain for the first time mainly in areas where inquilines have recently become abundant.
• 5 Since rates of parasitism of the gall former are still low, parasitoids are unlikely to play a major role in the population dynamics of this invading gall wasp at present, but the rapidly increasing inquiline and parasitoid attack could be a source of increased mortality for native cynipid species which are the alternative hosts of those parasitoid species.

     

Histological investigation on gall development induced by a worldwide invasive pest,Dryocosmus kuriphilus,on Castanea sativa

The Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae) is one of the most serious pests of chestnut trees worldwide. Wasps lay eggs into chestnut buds from mid Jane to late July, depending on latitude, and galls develop the following spring on new vegetation, reducing photosynthesis and nut production. We observed that modification of tissues surrounding D. kuriphilus eggs, which differentiate to form the larval chamber, started approximately 1 month after oviposition, shortly after eggs hatch. The larval chambers continued to increase slightly in size throughout the autumn months until January. After that, a period of stagnation, which corresponds to the plant's dormancy, occurred, followed by rapid growth from March to May, during the period from bud swelling to bud break. Galls continued to grow during the leaf expansion after bud break and stopped when plant organs achieved their final size. Our results have implications for the management of the pest, providing a better understanding of the critical time periods for the effective control.