Symbiotic associations between animals and microbes are widespread in nature, yet the factors controlling the abundance and distributions of particular symbionts are mostly unknown. Vertically transmitted (VT) symbionts can invade host populations by providing net benefits to hosts. While most beneficial symbiotic interactions that have been characterized are nutritional, other major pathways likely exist that facilitate symbiont transmission within host populations. Natural enemies, for example, are important selective forces in shaping the life history of many invertebrates and hosts may benefit from collaborations with microbes to aid in their defense.In this dissertation I have addressed the role of VT, facultative (= secondary) symbionts (SS) of Acyrthosiphon pisum in mediating interactions with an important natural enemy, the parasitic wasp, Aphidius ervi. I found that, in a common genetic background, two A. pisum SS (R- and T-type SS) confer resistance to A. ervi, by causing mortality to developing wasp larvae. Defensive mutualisms with microbes provide a mechanism for the spread and persistence of VT symbionts.A. pisum superinfected with both R- and T-type SS were found to be more resistant to parasitism that those singly infected with either SS. Despite this added benefit to resistance, R + T-type superinfections were rare in a survey of A. pisum symbionts, likely attributable to severe fecundity costs. R-type densities increased dramatically in superinfected hosts and over-proliferation of SS may result in poor aphid performance. Thus, interactions among the symbionts themselves also likely play a critical role in determining the distributions of symbionts in nature.I also found that four T-type isolates from A. pisum, and one from another aphid species, all conferred resistance to parasitism in the same A. pisum host background. The levels of resistance varied greatly among isolates, including one that conferred nearly complete resistance. A single T-type isolate was also found to confer similar levels of resistance in five A. pisum backgrounds. These results indicate that SS-mediated resistance is a general phenomenon in A. pisum and that the SS isolate is more important in determining the level of resistance than is the aphid genotype or interaction between isolate and aphid genotype.

Diachasmimorpha kraussii (Hymenoptera: Braconidae: Opiinae) is a koinobiont larval parasitoid of dacine fruit flies of the genus Bactrocera (Diptera: Tephritidae) in its native range (Australia, Papua New Guinea, Solomon Islands). The wasp is a potentially important control agent for pest fruit flies, having been considered for both classical and inundative biological control releases. I investigated the host searching, selection and utilisation mechanisms of the wasp against native host flies within its native range (Australia). Such studies are rare in opiine research where the majority of studies, because of the applied nature of the research, have been carried out using host flies and environments which are novel to the wasps. Diachasmimorpha kraussii oviposited equally into maggots of four fruit fly species, all of which coexist with the wasp in its native range (Australia), when tested in a choice trial using a uniform artificial diet media. While eggs laid into Bactrocera tryoni and B. jarvisi developed successfully through to adult wasps, eggs laid into B. cucumis and B. cacuminata were encapsulated. These results suggest that direct larval cues are not an important element in host selection by D. kraussii. Further exploring how D. kraussii locates suitable host larvae, I investigated the role of plant cues in host searching and selection. This was examined in a laboratory choice trial using uninfested fruit or fruit infested with either B. tryoni or B. jarvisi maggots. The results showed a consistent preference ranking among infested fruits by the wasp, with guava and peach most preferred, but with no response to uninfested fruits. Thus, it appears the wasp uses chemical cues emitted in response to fruit fly larval infestation for host location, but does not use cues from uninfested fruits. To further tease apart the role of (i) suitable and non-suitable maggots, (ii) infested and uninfested fruits of different plant species, and (iii) adult flies, in wasp host location and selection, I carried out a series of behavioural tests where I manipulated these attributes in a field cage. These trials confirmed that D. kraussii did not respond to cues in uninfested fruits, that there were consistent preferences by the wasps for different maggot infested fruits, that fruit preference did not vary depending on whether the maggots were physiologically suitable or not suitable for wasp offspring development, and finally, that adult flies appear to play a secondary role as indicators of larval infestation. To investigate wasp behaviour in an unrestrained environment, I concurrently observed diurnal foraging behaviours of both the wasp and one of its host fly in a small nectarine orchard. Wasp behaviour, both spatially and temporally, was not correlated with adult fruit fly behaviour or abundance. This study reinforced the point that infested fruit seems to be the primary cue used by foraging wasps. Wasp and fly feeding and mating was not observed in the orchard, implying these activities are occurring elsewhere. It is highly unlikely that these behaviours were happening within the orchard during the night as both insects are diurnal. As the final component of investigating host location, I carried out a habitat preference study for the wasp at the landscape scale. Using infested sentinel fruits, I tested the parasitism rate of B. tryoni in eucalyptus sclerophyll forest, rainforest and suburbia in South East Queensland. Although, rainforest is the likely endemic habitat of both B. tryoni and D. kraussii, B. tryoni abundance is significantly greater in suburban environments followed by eucalyptus sclerophyll forest. Parasitism rate was found to be higher in suburbia than in the eucalyptus sclerophyll forest, while no parasitism was recorded in the rainforest. This result suggests that wasps orient within the landscape towards areas of high host density and are not restricted by habitat types. Results from the different experiments suggest that host searching, selection and utilisation behaviour of D. kraussii are strongly influenced by cues associated with fruit fly larval feeding. Cues from uninfested fruits, the host larvae themselves, and the adult host flies play minimal roles. The discussion focuses on the fit of D. kraussii to Vinson’s classical parasitoid host location model and the implications of results for biological control, including recommendations for host and plant preference screening protocols and release regimes.

Insects have become increasingly resistant to chemical control methods, while at the same time public awareness of the harmful effects of synthetic pesticides has increased. The search for more environmentally 'friendly' means of pest suppression is gaining momentum and biological control (the use of natural enemies to reduce populations of noxious organisms) has become an increasingly sought-after option. Despite an increase in the establishment rate of insect natural enemies, classical biological control of arthropods is currently no more successful than it was one hundred years ago. Ecological theory relevant to population biology, food webs and diversity provides insight into how biological control agents can be made more effective, yet this theory has often been absent from the biological control literature. Examples of the use of ecological concepts (including intraguild predation, life-history omnivory and resource subsidies) in practical biological control are reviewed, and aspects of theory not yet considered in this context are discussed. Cereals are important as primary food crops, globally and within New Zealand. Possibly the greatest amount of damage sustained by cereal crops in New Zealand is caused by aphids and chemical control of these pests is very expensive relative to biological control. This thesis examines how biological control of the rose-grain aphid Metopolophium dirhodum (Walker) (Hemiptera: Aphidiidae) by the koinobiont, synovigenic endoparasitoid Aphidius rhopalosiphi De Stefani-Perez (Hymenoptera: Aphidiidae) can be enhanced by floral resource subsidies. The mechanisms underlying this enhancement were determined in a series of laboratory experiments and then tested in laboratory microcosms and in the field. Sugar resources significantly increased longevity and egg load in A. rhopalosiphi and another species of aphidiid, Diaeretiella rapae McIntosh. Pollen had no significant effect on longevity or egg load in these species. These results are discussed in terms of the effects of resource subsidies on egg- versus time-limitation. Laboratory microcosm experiments tested whether the mechanisms of increased potential fecundity via enhanced egg load and longevity translate into increased rates of parasitism (i.e., realised fecundity). Only treatments receiving sugar showed increased reproductive success. The presence of flowering buckwheat Fagopyrum esculentum Moench (c.v. Kitawase) (Polygonaceae) plants caused a slight, non-significant increase in rates of parasitism. Field surveys of natural aphid populations in a wheat Triticum aestivum (L.) (c.v. Otane) (Gramineae) field showed that proximity to floral buckwheat patches, distance to the nearest edge or the leeward end of the field were not significantly correlated with rates of parasitism. These variables were significantly correlated with aphid density in some surveys. Rates of parasitism were not correlated with aphid density. When aphid population density was controlled by experimental placement of aphids, proximity to floral resource patches significantly affected rates of parasitism. Parasitism rates were highest at the edges of buckwheat patches and declined exponentially with distance, eventually reaching zero beyond 14 m. Lower levels of parasitism were observed within the floral patches than at their edges. This phenomenon is compatible with the concept of spatio-temporal partitioning between parasitoid feeding and host-searching behaviours. Potential costs (increased predation risk, opportunity costs) and benefits (increased fecundity and longevity) associated with floral feeding by parasitoids, and temporal variation in these factors, are discussed. It is concluded that resource subsidies are not only effective in the practical enhancement of the efficacy of a specific biological control agent, but their use is based on a sound foundation in ecological theory that allows extension of these principles across taxa.

Hohenheim, Univ., Diss., 2006.

Parasitic wasps have a wide range of specific developmental interactions with their hosts due to a close "parasitoid-host" relationship and that allows the study of many ecological and evolutionary questions. In our work, we focused on the parasitoid-host interactions in the parasitic wasp Anaphes flavipes (Förster, 1841) (Hymenoptera: Mymaridae). Because A. flavipes show a high variability of morphology and size, they have been repeatedly described as a new species in the past. Thus, we first redescribed the species and we compared the variability of some morphological structures for wasps from different localities (Paper 1). The high variability of the body size of wasps is reviewed in the second article, where we propose that females of A. flavipes do not decide according to the generally known "trade-off" scheme, if they will have more smaller offspring or fewer bigger offspring. In fact, the number of offspring depends on the body size of the female ("body size-fitness" hypothesis), which determines the variable reproductive strategies of mothers (the number of individuals that are laid into one host by a single female) (Paper 2). The third manuscript deals with the reproductive strategy of gregarious parasitoids. The reproductive strategy does not have to be stable; in the case of A....

In this paper, 48 ichneumonid species of 14 genera, two tribes of the subfamily Pimplinae were listed on the basis of material collected during the time from 2003 to 2011 in four provinces from the Central Highlands of Vietnam and a lesser degree of previous records. Among 48 aforemention pimpline no less than 16 species were described recently as new for science. The diversity of topography and vegetation types resulted in the high level of species richness and new discoveries of the region. / Bài báo này giới thiệu danh sách 48 loài ong cự thuộc 14 giống, 2 tộc của phân họ Pimplinae dựa trên việc phân tích các mẫu vật thu tại Tây Nguyên trong thời gian từ năm 2003–2011 và một số ít loài dựa trên các ghi nhận trước đây. Trong số đó, 16 loài mới được mô tả trong thời gian 5 năm trở lại đây. Sự đa dạng về địa hình và các kiểu thảm thực vật khác nhau đã tạo nên tính đa dạng thành phần loài và các phát hiện mới của ong cự ở khu vực Tây Nguyên đầy tiềm năng này.

BACKGROUND:Numerous studies have investigated cospeciation between parasites and their hosts, but there have been few studies concerning parasitoids and insect hosts. The high diversity and host specialization observed in Anicetus species suggest that speciation and adaptive radiation might take place with species diversification in scale insect hosts. Here we examined the evolutionary history of the association between Anicetus species and their scale insect hosts via distance-based and tree-based methods.RESULTS:A total of 94 Anicetus individuals (nine parasitoid species) and 113 scale insect individuals (seven host species) from 14 provinces in China were collected in the present study. DNA sequence data from a mitochondrial gene (COI) and a nuclear ribosomal gene (28S D2 region) were used to reconstruct the phylogenies of Anicetus species and their hosts. The distance-based analysis showed a significant fit between Anicetus species and their hosts, but tree-based analyses suggested that this significant signal could be observed only when the cost of host-switching was high, indicating the presence of parasite sorting on related host species.CONCLUSIONS:This study, based on extensive rearing of parasitoids and species identification, provides strong evidence for a prevalence of sorting events and high host specificity in the genus Anicetus, offering insights into the diversification process of Anicetus species parasitizing scale insects.

Parasitoid wasps are hyperdiverse, with current estimates suggesting that they may account for up to 20% of all insect species. Though their ecological significance and their importance in integrated pest management cannot be denied, these taxa remain understudied and, due to their small size, are often overlooked. However, recent advances in molecular techniques are helping to reverse this trend by providing tools which scientists can use to better understand species limits and host interactions. Parasitoid wasps are often morphologically cryptic and their accurate delimitation requires the analysis of DNA sequence data from fast-evolving genes in addition to morphological characters. The research presented here demonstrates the utility of a new molecular locus in species delimitation. Also, a morphological key to the species of a genus occurring in America, north of Mexico is presented. The interactions between parasitoid wasps and their hosts are highly complex. On the wasp side, it involves the production venom, which likely contains bountiful natural resources. In this study, the venom proteins of wasps of the genus Megarhyssa (Hymenoptera: Ichneumonidae) are identified. Putative functions are assigned to these proteins and possible applications are discussed. One of the proteins identified is the enzyme, laccase, which is associated with the degradation and digestion of wood. The sequence of the gene coding for this laccase was analyzed and used to create recombinant proteins in a baculovirus-insect cell expression system. Future work investigating this enzyme is necessary to determine its activity against the plant cell wall. The research presented here provides insight into the identification and venom composition of ichneumonid wasps. The results contribute to our knowledge of this understudied taxon and indicate that there is much to be gained from further research in this field which will become increasingly practicable as molecular techniques advance and become more affordable.

The thesis is focused on phylogeny of the family Torymidae (Hymenoptera: Chalcidoidea) and evolution of their life-strategies. The study consists of general introduction to the phylogeny and classification of the family Torymidae chapter, four published papers in international journals and one manuscript prepared for submission. Firstly, our aim was to figure out the phylogenetic position of Torymidae as well as the position of other chalcidoid families inside superfamily Chalcidoidea (paper I and II). The supermatrix of sequencies of two ribosomal genes (18S rDNA and 28S rDNA) were developed for 649 species of chalcidoid taxa. However, family Torymidae was considered as polyphyletic group with the subfamily Megastigminae unrelated to the subfamily Toryminae (paper I). Monophyly of Torymidae was corroborated in another study (paper II) focused on molecular and morphological characters. We used a web-based, systematics workbench mx database for scoring 233 characters of 300 members of all chalcidoid families. Contrary to our previous only DNA-based study, we revealed also potential sister relationships of Torymidae with Ormyridae+Colotrechninae or Cerocephalinae+Diparinae respectively. Other paper (paper V) was focused on detailed study of Torymidae phylogeny. A total of 5 genes (18S rDNA, 28S rDNA,...

Toxoneuron nigriceps is a koinobiont larval endoparasitoid of the tobacco budworm, Heliothis virescens. No koinobiont parasitoid (which develop in active, growing hosts) has been reared completely in vitro. However, T. nigriceps has been partially reared in vitro, from egg stage through 2nd larval instar and from 2nd to 3rd larval instar. Despite reaching its final larval instar, T. nigriceps failed to develop past the larval stage. In the following study, two aspects of T. nigriceps development, post-egression tissue feeding and cocoon formation through pupation, were examined to determine why T. nigriceps failed to develop in vitro beyond the larval stage and how in vitro rearing might be improved to rear T. nigriceps to adulthood. The importance of post-egression feeding and the possibility of developing an artificial post-egression diet were examined by manipulating the post-egression feeding behavior of T. nigriceps. The tissues of pre- and post-tissue feeding T. nigriceps larvae were analyzed to investigate the macronutritional benefit of post-egression feeding. A preliminary artificial post-egression diet was tested, and the putative quality of T. nigriceps reared to adulthood using this diet was examined. The formation of subterranean pupation chambers by healthy and parasitized H. virescens larvae was examined to investigate potential behavioral manipulation by T. nigriceps during this process. Finally, the potential use of artificial pupation chambers by T. nigriceps was investigated. Post-egression feeding was determined to be a vital part of T. nigriceps development, contributing to both parasitoid size and survival to adulthood. A preliminary artificial post-egression diet was developed, which facilitated rearing of T. nigriceps from 3rd larval instar to adulthood. The dimensions and burial depth of chambers formed by parasitized H. virescens were significantly different from those of chambers formed by healthy H. virescens, suggesting that T. nigriceps manipulated host behavior during excavation. Toxoneuron nigriceps was able to use a number of common laboratory items as substitute pupation chambers during cocoon formation, and the merits of several artificial chambers were discussed. With provision of an adequate post-egression and artificial pupation chamber, in vitro rearing of T. nigriceps from the larval stage to adulthood may be possible.