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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.

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.

Trade-offs in resource allocation underlie key life history traits of organisms. My dissertation focuses on the size-number trade-off in clonal broods of offspring using the polyembryonic wasp, Copidosoma bakeri parasitizing immature stages of the moth Agrotis ipsilon. I aim to characterize responses of wasp brood size and individual body mass by manipulating the environments in order to understand the allocation pattern in the size number trade-off. In reviewing the functional forms of trade-off relationships in relation to resource constraints, I distinguish among three main trade-off types based on graphical representations of the relationship between the trade-off variables: linear, convex (inverse), and concave. The size-number trade-off in C. bakeri shows convex relationship. Characteristics of the trade-off are sex specific: female broods have larger body mass but smaller brood size than do male broods. When food intake of the host was increased, the trade-off between wasp body mass and brood size for both sexes shifts toward both higher wasp brood size and higher body mass. When the host has better access to food late in development, the size-number trade-off curve moves up and to the right on the graph. However, the trait combinations shift along the same trade-off curve toward greater wasp body mass but smaller brood size when the host development time is shorter due to more resources in early in development. I also investigate temperature effects on the size-number trade-off. C. bakeri brood size significantly increases with high temperature early in host development. There is no shift in the allocation pattern of the size-number trade-off with temperature. Finally, I test effects of body mass on longevity, fecundity, and mating competitiveness of C. bakeri. Larger body size increases female longevity, and mated females produce more eggs than unmated females. There are no significant relationships between male body mass and longevity or mating competitiveness. Mating reduces male longevity independent of body mass. The different impact of body mass on fitness between male and female wasps suggests the observed sex-specific allocation patterns of the size-number trade-off. Implications of the experiments and possible follow-up work are discussed.

Endosymbiosis is a pervasive phenomenon that has been a powerful force in insect evolution. In many well studied insect-bacterial associations, the bacteria can serve as reproductive manipulators, nutritional mutualists or defenders of their hosts. Fungi are also frequently associated with insects, and initial estimates suggest that these fungi are hyperdiverse. Saving a handful of examples, however, the functions of these fungi within insect hosts are largely unknown. This dissertation begins with a review that lays the conceptual groundwork for understanding bacterial and fungal endosymbiosis in insects. I make predictions about why one versus the other microbe might serve the insect, given any unique physiological, ecological or evolutionary conditions. I then aim to derive insights about microbial symbiosis by focusing on a particular system, that of brownbanded cockroaches, Supella longipalpa (Blattaria: Blattellidae) and their specialist wasp parasitoids, Comperia merceti (Hymenoptera: Encyrtidae). Here, I identify the symbiotic community of these two insects by using both culture-dependent and independent methods to characterize the vertically transmitted bacterial and fungal associates. Finally, I show that a heritable fungus in C. merceti, long presumed to be a mutualist, is parasitic under laboratory conditions: infected wasps incur fitness costs for housing the fungal symbiont relative to uninfected wasps. Additionally, although the fungus is not horizontally transmitted sexually, it is readily horizontally transmitted from the offspring of infected females to those of uninfected females that are using the same host.

Cotesia flavipes (Cameron) (Hym.: Braconidae), como outros cenobiontes, é capaz de regular seu hospedeiro, criando um ambiente que sustenta e promove o desenvolvimento de suas larvas, comumente em detrimento do hospedeiro. Substâncias derivadas do trato reprodutivo das fêmeas (proteínas ovarianas, veneno e polidnavírus) são injetadas no hospedeiro, afetando a resposta imune e outros processos fisiológicos com o propósito de regular os níveis hormonais, nutrição e comportamento. O presente trabalho teve por objetivo avaliar o papel dessas substâncias no crescimento e desenvolvimento de Diatraea saccharalis (Fabricius) (Lepidoptera: Crambidae), e avaliar como o parasitismo afeta a resposta imune do hospedeiro. Todas as substâncias derivadas da fêmea foram obtidas após a dissecação do parasitóide, através da coleta do reservatório de veneno ou dos ovários (proteínas ovarianas e polidnavírus) em tampão resfriado. As secreções foram processadas adequadamente e injetadas logo após a coleta. O veneno e as proteínas ovarianas + polidnavírus (PDV) foram injetados juntos ou separadamente em lagartas entre 0-12h do 6º instar. O efeito de cada um dos componentes isolados do parasitóide no desenvolvimento e crescimento do hospedeiro foi avaliado através de observações no ganho de peso, duração e viabilidade da fase larval e pupal. Os efeitos do parasitismo na resposta imune do hospedeiro foram avaliados tanto ao nível celular, através da contagem do número total de hemócitos e capacidade de encapsulação, como ao nível bioquímico, medindo-se a ativação da profenoloxidase e produção de óxido nítrico na hemolinfa das lagartas de D. saccharalis em diferentes estágios de desenvolvimento do parasitóide (0, 1, 3, 5, 7 e 9 dias após o parasitismo). As proteínas ovarianas do parasitóide e o PDV sozinho, ou co-injetado com o veneno, suspenderam o desenvolvimento larval do hospedeiro, enquanto que o veneno, sozinho, afetou o processo de metamorfose. A resposta imune do hospedeiro também foi afetada por C. flavipes, de maneira dependente do tempo. Lagartas parasitadas apresentaram declínio no número total de hemócitos a partir do 3º dia e a capacidade de encapsulação foi afetada ao longo do desenvolvimento do parasitóide. A atividade da fenoloxidase do hospedeiro foi alterada apenas no final do desenvolvimento imaturo do parasitóide, enquanto que o óxido nítrico foi afetado nas 24 h iniciais após parasitismo. / Cotesia flavipes (Cameron) (Hym., Braconidae), as other koinobionts, is capable of regulating the host development to produce an suitable host environment to sustain and promote its own larval development at the host expenses. Female-derived substances from the reproductive tract (ovarian proteins, venom, polydnavirus) are injected into the host, affecting the host immune response and other physiological processes aiming to regulate the host hormone levels, nutrition and behavior. Our goal was to evaluate the role of these substances on Diatraea saccharalis (F.) (Lepidoptera: Crambidae) growth and development, and how the parasitism affects the host immune response. All female-derived substances were collected after parasitoid dissection by collecting the venom reservoir or the ovaries (ovarian proteins and polydnavirus). Dissections were carried out in ice-cold buffer, collected tissues were processed accordingly and the desired substances injected immediately after collection. Venom and ovarian proteins+polydnavirus (PDV) were injected jointly and separated in 0-12 hold 6th instars of D. saccharalis. The effect of these substances on host development and growth was evaluated by measuring the host weight gain, larval and pupal survivorship and developmental time. The effects of the parasitism on the host immune response was evaluated either at the cellular level, by measuring the total hemocyte count and the encapsulation capacity, and at the biochemical level, by measuring the prophenoloxidase activity and nitric oxide levels at different stages of parasitoid development (0, 1, 3, 5, 7 and 9 days after parasitism). Parasitoid ovarian proteins and PDV alone or co-injected with the venom arrested the host larval development, while the venom by itself only affected the host metamorphosis process. The host immune response was also affected by C. flavipes at a time-dependent manner. The total hemocyte count dropped at day 3 of parasitism, while the host encapsulation capacity was reduced during parasitoid development. The host prophenoloxidase activity was also affected mainly towards the end of parasitoid larval development, while the nitric oxide at the first 24 h after parasitism.

Phytophagous (plant-feeding) insects are extremely species-rich and typically display tight host associations (meeting and mating on or near their host plant) with one or a small number of hosts. This specialized lifestyle can promote diversification through assortative mating, ultimately leading to genetically differentiated host races (host associated differentiation; HAD). It has been shown that HAD can cascade up to the parasitic wasps (parasitoids) that utilize the phytophagous insects as hosts. Cascading HAD occurs when there is genetic differentiation among parasitoids as a result of differential host plant use by their host insects. Thus, host switching can promote parasitoid diversification as well. Here, I present three studies designed to help understand aspects of parasitoid shifts to novel hosts and environments. All of the studies in this dissertation utilize the Rhagoletis complex of flies and their associated parasitoids. Specifically, I address i) the role of subtle trait variation and environmental context in predicting successful parasitoid host shifts; ii) whether parasitoid host discrimination (a trait that can influence host shifts) is an innate or learned behavior; and iii) whether contemporary patterns of host shifts among parasitoids are echoed by historical host shifts in cophylogenetic analyses of host and parasitoid genera? Towards my first aim, I present a phenomenological model developed to predict successful host shifts by parasitoids. The simulations of the model explore how environmentally mediated traits can affect successful parasitoid colonization of a new host. For my second aim, I hypothesize that behaviors impacting parasitoid host plant preferences host shifts will be genetically based rather than a learned behavior. Shifting to a new host plant has been shown to cause reproductive isolation in phytophagous insects because of strong fidelity with their host plant. Parasitoids, however, have no direct contact with the host plant as they develop entirely within the host insect. The differences in life history traits could result parasitoid host shifts being driven by random changes in host preferences. I present preliminary results suggesting that parasitoids preferentially respond to their ancestral host plant’s olfactory cues, suggesting that host preferences have a genetic basis. Finally, I present a cophylogenetic analysis of Rhagoletis hosts and their parasitoids. I find that cospeciation is the most common coevolutionary event, although there is evidence of recent host shifting that contributes to current parasitoid species diversity. The results of these studies can help us understand how host shifts can act as a potential mechanism driving diversification in parasitoids.

Invasive insects are considered one of the most serious threats affecting forests today; however, surprisingly little research has addressed the impacts of invasive species establishment on native forest insect communities. Such information is lacking for even the most thoroughly studied invasive forest insect, the gypsy moth. Using gypsy moth as a case study, my thesis addresses the questions: What are the ecological impacts of an exotic forest insect upon its establishment in a new community of native species? Does the community shift after the invasive establishes, and if so, what are the drivers in this realignment? I used multivariate analysis to assess native caterpillar communities collected in forest stands with and without a history of gypsy moth outbreak. I found that gypsy moth outbreak history had no significant effects on native caterpillar communities; however, current gypsy moth abundance was related to shifts in the structure of late season caterpillar assemblages. These results suggest that gypsy moth may affect native caterpillar communities through short-term mechanisms but not through long-term ecological changes. I used quantitative food webs to investigate the effects of gypsy moth on native host-parasitoid webs from the same caterpillar communities, and found that food web structure was resilient to both gypsy moth outbreak history and current abundance. The gypsy moth shared few parasitoids with native species in my study sites, none of numerical significance, thus minimizing the opportunity for enemy-mediated indirect interactions. Finally, I conducted a greenhouse experiment and found that early spring feeding by forest tent caterpillar can indirectly influence gypsy moth susceptibility to its virus, demonstrating that the complex interactions that can occur between native and exotic species do not always benefit the invader. Overall, I argue that the establishment of the gypsy moth into North American forests will not cause major changes in native caterpillar communities.

Invasive insects are considered one of the most serious threats affecting forests today; however, surprisingly little research has addressed the impacts of invasive species establishment on native forest insect communities. Such information is lacking for even the most thoroughly studied invasive forest insect, the gypsy moth. Using gypsy moth as a case study, my thesis addresses the questions: What are the ecological impacts of an exotic forest insect upon its establishment in a new community of native species? Does the community shift after the invasive establishes, and if so, what are the drivers in this realignment? I used multivariate analysis to assess native caterpillar communities collected in forest stands with and without a history of gypsy moth outbreak. I found that gypsy moth outbreak history had no significant effects on native caterpillar communities; however, current gypsy moth abundance was related to shifts in the structure of late season caterpillar assemblages. These results suggest that gypsy moth may affect native caterpillar communities through short-term mechanisms but not through long-term ecological changes. I used quantitative food webs to investigate the effects of gypsy moth on native host-parasitoid webs from the same caterpillar communities, and found that food web structure was resilient to both gypsy moth outbreak history and current abundance. The gypsy moth shared few parasitoids with native species in my study sites, none of numerical significance, thus minimizing the opportunity for enemy-mediated indirect interactions. Finally, I conducted a greenhouse experiment and found that early spring feeding by forest tent caterpillar can indirectly influence gypsy moth susceptibility to its virus, demonstrating that the complex interactions that can occur between native and exotic species do not always benefit the invader. Overall, I argue that the establishment of the gypsy moth into North American forests will not cause major changes in native caterpillar communities.

Global warming and increasing atmospheric nitrogen deposition are ranked as second and third most important global drivers of biodiversity loss. Widespread species losses have deep implications for the functioning of ecosystems, the delivery of essential ecosystem services and their resilience to future environmental perturbations. There is growing recognition that interactions between species play a crucial role in determining the response of ecosystems to global environmental changes. Moreover, evidence of synergistic effects between global change drivers has prompted numerous calls to integrate multiple drivers in ecological research. Nevertheless, empirical studies assessing the impacts of temperature and nitrogen on communities at multiple trophic levels are largely absent. This thesis explores the effects of temperature and nitrogen on a tri-trophic system comprising plants, herbivores and natural enemies. The first chapter shows impacts of the drivers on the composition and phenology of an herbivore community. The second chapter highlights changes in biomass under the treatments at three trophic levels. The third chapter explores, for the first time, the impacts of temperature and nitrogen on quantitative food webs. Finally, the last data chapter uses body size as an important species trait to gain insights on the mechanisms causing shifts in food web structure. The key findings of this thesis were i) trophic interactions largely mediated the effects of both global change drivers ii) In particular, strong bottom-up effects determined the system response, with herbivores responding positively and consistently more so than plants and parasitoids in particular. However, iii) this contrasting response was not explained by a phenological mismatch. iv) Food-web structure responded to the changes in composition of herbivores and parasitoids, but shifts in interaction structure did not affect the resilience of the food. However, temperature and nitrogen impacted host-parasitoid food-web structure by altering the response of parasitoid species to host density and size structuring, which is likely to bear consequences on host-parasitoid co-evolution and future food-web architecture and stability. Finally, v) we found frequent, non-additive interactions between the global change drivers. We conclude that co-occurring temperature and nitrogen are likely to alter food-web structure and overall ecosystem balance, with increasing herbivore dominance likely to have important implications for ecosystem functioning and food-web persistence.