Species Diversity in the Parasitoid Genus Asobara (Hymenoptera: Braconidae) from the Native Area of the Fruit Fly Pest Drosophila suzukii (Diptera: Drosophilidae)

Drosophila suzukii (Matsumura), commonly known as Spotted Wing Drosophila (SWD), is a worldwide serious economic threat to the production of berries and stone fruits. The chemical control widely used against this pest is often not able to preventing yield losses because wild flora offers an abundance of fruits to D. suzukii where the pest is able to reproduce and from where it recolonizes neighbouring cultivated fields. Alternatively, within Integrated Pest Management protocols for D. suzukii, biological control could play a key role by reducing its populations particularly in non-cultivated habitats, thus increasing the effectiveness and reducing the side negative effects of other management strategies. Because of the scarcity and of the low efficiency of autochthonous parasitoids in the new invaded territories, in the last few years, a number of surveys started in the native area of D. suzukii to find parasitoid species to be evaluated in quarantine structures and eventually released in the field, following a classical biological control approach. This paper reports the results of these surveys carried out in South Korea and for the first time in China. Among the parasitoids collected, those belonging to the genus Asobara Foerster resulted dominant both by number and species diversity. By combining morphological characters and the mitochondrial COI gene as a molecular marker, we identified seven species of Asobara, of which two associated with D. suzukii, namely A. japonica and A leveri, and five new to science, namely Asobara brevicauda, A. elongata, A mesocauda, A unicolorata, A. triangulata. Our findings offer new opportunity to find effective parasitoids to be introduced in classical biological control programmes in the territories recently invaded by D. suzukii.     

Host range of Asobara japonica (Hymenoptera: Braconidae), a larval parasitoid of drosophilid flies

We studied the host range of Asobara japonica, a larval‐pupal parasitoid of drosophilid flies. Habitat selection was found to be an important determinant of host range in this parasitoid; it attacked drosophilid larvae breeding on banana and mushrooms, but seldom attacked those breeding on decayed leaves. This parasitoid was able to use diverse drosophilid taxa as hosts. Attack by A. japonica sometimes killed hosts at the larval stage, and therefore parasitoid larvae also died. Drosophila elegans and D. busckii suffered particularly high larval mortality due to the attack by A. japonica (in the latter species only when young larvae were attacked). Many individuals of D. subpulchrella also died at the pupal stage without producing parasitoids when they were parasitized at the late larval stage. In contrast, D. bipectinata, D. ficusphila, D. immigrans, D. formosana and D. albomicans were resistant to attack: large proportions of the larvae of these drosophilid species grew to adulthood, even in the presence of parasitoids. On the basis of phylogenetic information, we concluded that phylogenetic position has only limited importance as a factor determining whether a species is suitable as a host for A. japonica, at least within the genus Drosophila.     

Physiological consequences of immune response by Drosophila melanogaster (Diptera: Drosophilidae) against the parasitoid Asobara tabida (Hymenoptera: Braconidae)

Abstract Parasites can exert a wide range of negative effects on their hosts. Consequently, hosts that can resist infection should have a selective advantage over nonresistant conspecifics. Yet, host populations remain susceptible to some parasites. Could genetic heterogeneity in the host's ability to resist parasites reflect costs of mounting an immune response? Previous work on Drosophila melanogaster establishes that maintaining the ability to mount an immune response decreases larval competitive ability. Moreover, mounting an immune response decreases fitness. I report on the impact of mounting an immune response on fitness of D. melanogaster survived parasitism by Asobara tabida. I used isofemale lines to determine whether genotype influences the costs of immune response. I examined fitness consequences both to larvae and adults. Survivors of parasitism show no measurable decrease in larval fitness (development time) but have decreased adult fitness (population growth rates), probably because of their smaller size.     

The effect of four fruit species on the parasitization rate of Anastrepha fraterculus (Diptera: Tephritidae,Trypetinae) by Diachasmimorpha longicaudata (Hymenoptera: Braconidae,Opiinae) under laboratory rearing conditions

In the laboratory, the effect of host fruit species Citrus paradisi, C. aurantium, Prunus persica, and Psidium guajava on A. fraterculus parasitism by Diachasmimorpha longicaudata was studied. The number of ovipositor-probing events and the probing-time were documented to evaluate the role of fruit chemistry, and epicarp and mesocarp thickness, respectively. The relationship between the parasitization rate and fruit size in particular plant species was analyzed by applying a simple regression. Results showed that guava and peach yielded significantly more parasitoids than both Citrus spp. Probably, the parasitization rate of D. longicaudata on A. fraterculus would be influenced in part by chemical and physical factors from fruit species.     

Phylogeny of a paradigm lineage: the Drosophila melanogaster species group (Diptera: Drosophilidae)

Although Drosophila melanogaster is a paradigm eukaryote for biology, relationships of this species and the other 174 species in the melanogaster species group are poorly explored and ambiguous. Gene regions of Cytochrome oxidase II (mt:CoII ), Alcohol dehydrogenase ( Adh ) and hunchback ( hb ) were sequenced and analysed phylogenetically to test prior hypotheses of relationships for the group based on chromosomes, morphology, and 28S rRNA gene sequences. A simultaneous cladistic analysis of the three newly sequenced gene regions produced a single well-resolved phylogeny for 49 exemplar species representing eight subgroups. Monophyly of each of the ananassae , melanogaster , montium , and takahashii subgroups is supported; the suzukii subgroup is polyphyletic. This phylogeny is consistent with variation in significant morphological structures, such as the male sex comb on the fore tarsus. The broad range of morphological variation among these species is interpreted and the applicability to evolution and developmental investigations is discussed. This phylogeny facilitates comparative investigations, such as gene family evolution, transposable element transmission, and evolution of morphological structures. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 21–37.     

Seasonal monitoring of Drosophila suzukii (Diptera: Drosophilidae) in a mixed fruit production system

Drosophila suzukii Matsumura (Diptera: Drosophilidae), an invasive pest native to Southeast Asia, is now reported throughout North America and Europe. We used traps baited with apple cider vinegar to monitor D. suzukii adult presence in multiple crops and associated fruiting plants at the Wolfskill USDA Germplasm Repository in Winters, CA, USA from 2011 to 2013. Traps were placed in small (~ 160 m × 40 m on average) almond, apricot, cherry, fig, grape, mulberry, peach, persimmon, plum, and pomegranate deciduous fruit orchard blocks as well as a citrus block and evergreen trees located near a house at the repository. D. suzukii was present in all blocks with the greatest monthly deciduous fruit captures in the cherry and fig blocks. Few D. suzukii were captured in almond, apricot, pomegranate and grape blocks. Deciduous fruit blocks had two distinct periods of trap capture: spring through midsummer and again in fall. Most deciduous fruit blocks had low trap captures during the hottest summer months (August to September) and the coldest winter months (December to April). However, from late December through mid-January, high trap captures were associated with the citrus and house sites. This study provides seasonal trapping data of D. suzukii adults in an unsprayed multi-crop mosaic, and may serve as a model of adult capture patterns across smaller mixed-crop commercial orchards and associated urban landscapes.     

The relative fitness of Drosophila melanogaster (Diptera,Drosophilidae) that have successfully defended themselves against the parasitoid Asobara tabida (Hymenoptera,Braconidae)

Drosophila melanogaster larvae defend themselves against parasitoid attack via the process of encapsulation. However, flies that successfully defend themselves have reduced fitness as adults. Adults which carry an encapsulated parasitoid egg are smaller and females produce significantly fewer eggs than controls. Capsule-bearing males allowed repeated copulations with females do not show a reduction in their number of offspring, but those allowed to copulate only once did. No differences were found in time to first oviposition in females, or in time to first copulation in males. We interpret the results as arising from a trade-off between investing resources in factors promoting fecundity and mating success, and in defence against parasitism. The outcome of this investment decision influences the strength of selection for defence against parasitism.     

Parasitoids from Azores (Hymenoptera: Encyrtidae,Pteromalidae, Braconidae): potential use in integrated pest management against Ceratitis capitata (Diptera: Tephritidae)

Two hymenopterous parasitoids, Halticoptera patellana and Toxeumorpha nigricola, were recovered from Mediterranean fruit fly pupae on São Miguel Island, Azores. Both are new records for the Azores, and as Medfly parasitoids. Field-collected Tachinaephagus zealandicus was maintained on Medfly for 10 generations, but if it has potential as a biocontrol agent remains an open question.     

Susceptibility of cold hardy grapes to Drosophila suzukii (Diptera: Drosophilidae)

Drosophila suzukii (Diptera: Drosophilidae), known commonly as spotted wing drosophila, is a vinegar fly originating from South‐East Asia and a major pest to many soft‐skinned fruits. Due to the species recent arrival in North America in 2008, many fruit varieties are yet untested for susceptibility to infestation. While previous work has focused on Vitis vinifera, this study aimed to determine grape susceptibility of cold hardy varieties based on hybrids of V. labrusca, V. riparia and V. vinifera. Field sampling was conducted in Southern Wisconsin (USA) vineyards to establish adult and larval abundance and determine whether the number of adults caught in traps correlates with fruit infestation. Host susceptibility was further assessed through no‐choice bioassays of both intact and damaged fruits. The field study found D. suzukii adults present in all varieties, low larval abundance and no correlation between adult abundance and larval presence. Peak adult abundance occurred mid‐season between veraison and harvest, while larval infestation rates were highest near harvest. In laboratory no‐choice tests, significantly more eggs, larvae and adults occurred in damaged than undamaged grapes. In damaged grapes, larvae and adult abundance was comparable between varieties and to the highly susceptible control of undamaged raspberry; however, D. suzukii developed significantly faster in raspberry than grapes. Fruit characteristics (°Brix, titratable acidity, pH) in grapes were uncorrelated with D. suzukii performance. Together, these findings suggest that cold hardy grapes are overall resistant to D. suzukii if intact and highly susceptible if damaged.     

Drosophila subpulchrella, a new species of the Drosophila suzukii species subgroup from Japan and China (Diptera: Drosophilidae)

Drosophila (Sophophora) subpulchrella Takamori and Watabe, sp. nov., of the D. suzukii subgroup in the D. melanogaster species group, is described from Japan and southern China, and compared with its sibling species, D. pulchrella Tan et al. distributed in the Yun‐Gui Highland, south‐western China. The results of cross‐experiments show a complete pre‐mating isolation between D. subpulchrella and D. pulchrella.     

Response of Drosophila suzukii (Diptera: Drosophilidae) to extreme heat and dryness

相似文献   

Humidity affects populations of Drosophila suzukii (Diptera: Drosophilidae) in blueberry

Temperature and humidity affect insect physiology, survival, fecundity, reproductive status and behaviour. Complementing previous work investigating the effects of temperature on adult survival and fecundity of the invasive frugivorous pest, Drosophila suzukii (Matsumura), this study was conducted to determine the effect of humidity on D. suzukii larval development, adult survival, fecundity and reproductive status using blueberry as a host substrate. The five constant humidity levels in laboratory bioassays were 20, 33, 71, 82 and 94% RH at 20.6 ± 0.2°C. As RH increased, fecundity and longevity increased. At the higher humidity levels, RH had limited impact on mean generation times (T), larval development and eclosion times. The highest net reproductive rate (R_o = 68) and highest intrinsic rate of population increase (r_m = 0.17) were both recorded at 94% RH. The reproductive status of females, as indicated by the number of mature oocytes per female, was significantly greater at 82 and 94% RH, compared to 71% RH. In addition to the laboratory procedures, we correlated field trap captures over an 81‐day summer period to relative humidity (RH) levels in close proximity to those traps. In the field, low ambient humidity levels resulted in decreased trap captures. A humidity‐dependent population model predicted lower densities of D. suzukii relative to populations at higher humidity. This study supports the hypothesis that cultural practices that minimize lower humidity levels in crops can contribute to the management of D. suzukii. Such methods may include open pruning, drip irrigation and field floor management.     

Acceptability and suitability of six fruit fly species (Diptera: Tephritidae) for Kenyan strains of Psyttalia concolor (Hymenoptera: Braconidae)

Host acceptability and suitability Psyttalia concolor (Szépligeti) is a koinobiont, larval parasitoid of tephritid fruit flies. Individuals of P. concolor were field-collected from coffee in the central highlands of Kenya, and cultured initially on Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann). They were then examined for their ability to oviposit in and develop on five other tephritid species that are pests in Kenya. In addition to the medfly, acceptability for oviposition and suitability for development were tested against the mango fruit fly, Ceratitis cosyra (Walker), the Natal fruit fly, Ceratitis rosa Karsch, Ceratitis fasciventris (Bezzi), Ceratitis anonae Graham and the melon fruit fly, Bactrocera cucurbitae (Coquillett). Ceratitis capitata and C. cosyra were accepted as hosts significantly more often than the other species. Superparasitism was recorded only from C. capitata and C. cosyra. Two days after oviposition, parasitoid eggs in C. fasciventris and B. cucurbitae were encapsulated, whereas those in C. rosa and C. anonae were encapsulated, and often melanized. Ceratitis capitata was the most suitable host for Kenyan populations of Psyttalia concolor in terms of progeny production, and proportion of female progeny.     

Associations of yeasts with spotted-wing Drosophila (Drosophila suzukii; Diptera: Drosophilidae) in cherries and raspberries

A rich history of investigation documents various Drosophila-yeast mutualisms, suggesting that Drosophila suzukii similarly has an association with a specific yeast species or community. To discover candidate yeast species, yeasts were isolated from larval frass, adult midguts, and fruit hosts of D. suzukii. Terminal restriction fragment length polymorphism (TRFLP) technology and decimal dilution plating were used to identify and determine the relative abundance of yeast species present in fruit juice samples that were either infested with D. suzukii or not infested. Yeasts were less abundant in uninfested than infested samples. A total of 126 independent yeast isolates were cultivated from frass, midguts, and fruit hosts of D. suzukii, representing 28 species of yeasts, with Hanseniaspora uvarum predominating. This suggests an association between D. suzukii and H. uvarum that could be utilized for pest management of the highly pestiferous D. suzukii.     

An ultrastructural and molecular study of Tubulinosema kingi Kramer (Microsporidia: Tubulinosematidae) from Drosophila melanogaster (Diptera: Drosophilidae) and its parasitoid Asobara tabida (Hymenoptera: Braconidae)

Tubulinosema kingi is a pathogen of Drosophila spp. that was originally described 40 years ago. Although Drosophila melanogaster is widely used as a model organism for biological research, only limited data about microsporidia infecting Drosophila have been published so far and very little is known about the ultrastructure of T. kingi. In this study, we present the results of ultrastructural and molecular examinations of T. kingi. The whole life cycle took place in direct contact with the host cell cytoplasm and all examined life cycle stages contained a diplokaryon. Very few membrane elements were present in early merogonial stages, but their number and order of arrangement increased as the life cycle proceeded. The cell membrane of meronts had a surface coat of tubular elements that encircled the cell. Later, numerous electron-dense strands without any ornamentation accumulated on the plasma membrane, indicating that cells had entered sporogony. The cell membrane of sporonts was covered by electron-dense material. The polar filament in the spores was slightly anisofilar with the last three or four coils being smaller in diameter. The polar filament has 10 to 14 coils which were arranged predominantly in a single row, but in many spores, one winding of the coiled polar filament was located inside the outer coils. In some spores, the polar filament was irregularly arranged in two or even three rows. Molecular analysis showed that all Tubulinosema spp. are closely related and form a clade of their own that is distinct from the Nosema/Vairimorpha clade. All these ultrastructural and molecular features are in concordance with the family Tubulinosematidae and the genus Tubulinosema which reinforces the recent reclassification of this microsporidium.     

Preference and performance of Diachasmimorpha kraussii (Fullaway) (Hymenoptera: Braconidae) on five commercial fruit species

Diachasmimorpha kraussii is a polyphagous endoparasitoid of dacine fruit flies. The fruit fly hosts of D. krausii, in turn, attack a wide range of fruits and vegetables. The role that fruits play in host selection behaviour of D. kraussii has not been previously investigated. This study examines fruit preference of D. kraussii through a laboratory choice‐test trial and field fruit sampling. In the laboratory trial, oviposition preference and offspring performance measures (sex ratio, developmental time, body length, hind tibial length) of D. kraussii were investigated with respect to five fruit species [Psidium guajava L. (guava), Prunis persica L. (peach), Malus domestica Borkh. (apple), Pyrus communis L. (pear) and Citrus sinensis L. (orange)], and two fruit fly species (Bactrocera jarvisi and B. tryoni). Diachasmimorpha kraussii responded to infested fruit of all fruit types in both choice and no‐choice tests, but showed stronger preference for guava and peach in the choice tests irrespective of the species of fly larvae within the fruit. The wasp did not respond to uninfested fruit. The offspring performance measures differed in a non‐consistent fashion between the fruit types, but generally wasp offspring performed better in guava, peach and orange. The offspring sex ratio, except for one fruit/fly combination (B. jarvisi in apple), was always female biased. The combined results suggest that of the five fruits tested, guava and peach are the best fruit substrates for D. krausii. Field sampling indicated a non‐random use of available, fruit fly infested fruit by D. kraussii. Fruit fly maggots within two fruit species, Plachonia careya and Terminalia cattappa, had disproportionately higher levels of D. krausii parasitism than would be expected based on the proportion of different infested fruit species sampled, or levels of fruit fly infestation within those fruit.     

Fitness of two sibling species of Asobara (Braconidae:Alysiinae), larval parasitoids of Drosophilidae in different microhabitats

Abstract. 1. Two sibling species of larval endoparasitoids of Drosophilidae: Asobara tabida (Nees) and A.rufescens (Foerster) occur in the same macrohabitat, but inhabit different microhabitats. Each species is most attracted by odours of its own microhabitat.
2. In order to assess the adaptive value of the microhabitat preference we studied the survival of both parasitoids in the major host species that occur in these microhabitats.
3. Survival in the major host in the preferred microhabitat was shown to be 40% higher for A.tabida and 30% higher for A.rufescens when compared to survival in the major host in the non-preferred microhabitat.
4. Measurements of developmental rates, specific mortalities and dry weights of the parasitoids suggest that the differential survival is due to differences in synchronization with the hosts.
5. The possible evolutionary consequences of some biological characteristics in Asobara are discussed. Microhabitat selection, differential survival, development and mating behaviour are attributes likely to have played a role in the reduction of gene flow between populations of the ancestral species, either in primary or in secondary sympatry.     

Rearing of the fruit fly parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae) on X-ray irradiated larvae of Ceratitis capitata (Diptera: Tephritidae)

We evaluated the effects of X-ray irradiation on larvae of Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), with the aim of finding a treatment that prevented adult fly emergence, yet did not adversely affect larval quality as rearing hosts for the parasitoid Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Ichneumonidae). Two experiments were carried out. In the first experiment, small numbers of larvae were held in Petri dishes with and without small amounts of rearing media and then irradiated with doses of X-rays ranging from 0 to 8333.6 R (equivalent to 80 Gy). In the second experiment, higher numbers of larvae were held in a manner resembling mass-rearing conditions, and were then irradiated with X-rays ranging from 0 to 10,417 R (equivalent to 100 Gy). In both experiments, the only factor that significantly affected fly emergence was irradiation of larvae. Fly emergence decreased markedly as the irradiation dose increased, and complete suppression of fly emergence was achieved at 6250.2 R (equivalent to 60 Gy) when larvae were irradiated in small batches with or without rearing media. Irradiation also affected the fertility of those flies that did emerge following treatment. In the second experiment, we found the parasitoids reared from irradiated larvae produced a higher parasitism rate and a higher number of female offspring than did parasitoids reared from control (non-irradiated) larvae. Mean fecundity of F1 parasitoids reared from irradiated larvae were affected positively by irradiation only at the 8333.6 R (80 Gy) dose. Our results show that X-ray irradiation can be used to inhibit fruit fly adult emergence and that irradiated larvae are at least as good a rearing substrate as non-irradiated larvae. Future studies should focus on the adjustment of our findings to a mass-rearing scale.     

Physiological status of Drosophila suzukii (Diptera: Drosophilidae) affects their response to attractive odours

An improved understanding of the biology of the invasive pest, Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), is critical for the development of effective management strategies. Trapping is one technique used for both detection and control; however, the efficacy of trapping can vary depending on the target insect's physiological state, its behavioural priorities and the type of attractant used in the trap. We conducted a series of caged trapping experiments and a greenhouse trapping experiment to investigate the effects of D. suzukii feeding status, age, mating status, ovipositional status and seasonal morph type on the capture rate of traps baited with fermentation odours. Starved flies were trapped at greater rates compared to fed flies; more virgin flies were trapped than mated flies; flies deprived of an oviposition substrate were trapped more frequently than flies given an oviposition substrate. It is still unclear whether age or seasonal morphology affect bait response. Lastly, a caged choice experiment investigated the relationship between female reproductive status and attraction to fermentation or fruit odours. Fermentation‐based traps captured female flies regardless of their reproductive status but, ripe fruit‐based traps were more attractive to flies with more than seven eggs. In summary, studies that use fermentation‐based traps should recognize that capture rates of D. suzukii will depend on the feeding, mating and oviposition experiences of the population; also, fruit‐based traps may better target gravid females.     

白蜡吉丁柄腹茧蜂的寄生和繁殖生物学

白蜡吉丁柄腹茧蜂Spathius agrili Yang是我国天津地区寄生白蜡窄吉丁幼虫的一种重要天敌。本研究通过林间调查并结合室内观察,测定了该寄生蜂在不同营养条件下的存活时间和不同时期的产卵量,调查了不同寄主大小对寄生蜂后代生长发育的影响,以及寄生蜂对不同深度树皮下的寄主利用情况。结果表明：不同营养条件下白蜡吉丁柄腹茧蜂的存活时间差异很大,补充20％蜂蜜水的成蜂存活时间明显延长,最长可达3个月。相同条件下雌蜂的存活时间比雄蜂略长。在补充营养的条件下,茧蜂的产卵时间最长可达8周,产卵的高蜂在羽化后第2周前后,平均每雌每周产卵量在整个产卵期内基本保持在8～12粒上下,没有大的波动。茧蜂低龄幼虫也可能分泌毒素主动攻击寄主,提高了寄生蜂后代的生存能力。寄主个体大,营养量也大,寄生其上的茧蜂幼虫成功完成发育的数量随之增加,表明寄主营养资源的多少直接影响到寄生蜂后代的生长发育和存活。白蜡吉丁柄腹茧蜂对1～6.5 mm树皮深度范围内的适龄寄主幼虫均能够寄生利用,约92％的被寄生白蜡窄吉丁幼虫个体分布于1～4 mm深度的树皮下,在3 mm厚的树皮下分布的数量最多。茧蜂数量与树皮深度大致上呈抛物线关系,理论上在树皮深度为3.97 mm处茧蜂分布数量最多。