O objetivo do presente trabalho foi avaliar, em laboratório, a seletividade de produtos químicos, utilizados na citricultura, ao parasitóide de ovos Trichogramma atopovirilia Oatman & Platner (Hymenoptera: Trichogrammatidae), inimigo natural com maior potencial de uso para o controle biológico de Gymnandrosoma aurantianum Lima (Lepidoptera: Tortricidae), uma das principais pragas dos citros. Os ensaios foram conduzidos seguindo as normas estabelecidas pela IOBC/WPRS para a avaliação dos efeitos nocivos de agroquímicos aos inimigos naturais de insetos. Utilizou-se água destilada como testemunha e foram selecionados cinco inseticidas (imidacloprid, deltametrina, Bacillus thuringiensis, óleo mineral e triflumuron), três acaricidas (abamectina, dicofol e espirodiclofeno) e quatro fungicidas (oxicloreto de cobre, piraclostrobina, difenoconazole e mancozebe) utilizados para o controle do bicho-furão-dos-citros e outras pragas e doenças de citros. Foram realizados ensaios relativos à exposição de adultos, forma mais suscetível do parasitóide, aos produtos; contato direto com formas imaturas (ovo-larva, pré-pupa e pupa) no interior dos ovos do hospedeiro natural G. aurantianum e do hospedeiro alternativo, Anagasta kuehniella e aceitação, pelas fêmeas do parasitóide, de ovos tratados com os produtos. Posteriormente, os produtos foram classificados, conforme a redução causada no parasitismo ou porcentagem de emergência em: 1, inócuo (<30%); 2, levemente nocivo (30 a 79%); 3, moderadamente nocivo (80 a 99%); 4, nocivo (>99%). Os inseticidas, acaricidas e fungicidas testados afetaram o parasitismo de T. atopovirilia. Não houve diferença no comportamento de T. atopovirilia fosse ele criado no hospedeiro natural ou no alternativo. Os produtos menos seletivos aos adultos do parasitóide foram deltametrina, abamectina, óleo mineral e imidacloprid. Em relação aos estágios imaturos do parasitóide, todos os produtos foram inócuos, exceto óleo mineral, que provocou redução de 100% na porcentagem de emergência; abamectina, triflumuron, imidacloprid foram levemente prejudiciais ao período ovo-larva. Abamectina e triflumuron tiveram maior efeito sobre a capacidade de parasitismo dos indivíduos emergidos quando o tratamento ocorreu durante a fase de pupa do parasitóide. Os produtos Bacillus thuringiensis, difenoconazole, oxicloreto de cobre, piraclostrobina, dicofol, espirodiclofeno, considerados inócuos ou levemente prejudiciais em todas as etapas dos ensaios em laboratório, podem ser utilizados em programas de Manejo Integrado de Pragas, em associação com o controle biológico, sem prejuízo para o parasitóide e contribuindo para a sustentabilidade da citricultura. / This study sought to evaluate in the laboratory the selectivity of chemical products used in citriculture to the egg parasitoid Trichogramma atopovirilia Oatman & Platner (Hymenoptera: Trichogrammatidae), the natural enemy with the greatest potential for use in the biological control of Gymnandrosoma aurantianum Lima (Lepidoptera: Tortricidae), one of the main pests of citrus. Assays were conducted following the standards established by IOBC/WPRS to evaluate the harmful effects of agrochemicals to natural enemies of insects. Distilled water was used as a control and five insecticides (imidacloprid, deltamethrin, Bacillus thuringiensis, mineral oil, and triflumuron), three miticides (abamectin, dicofol, and spirodiclofen), and four fungicides (copper oxychloride, pyraclostrobin, difenoconazole, and mancozeb) used to control the citrus fruit borer and other pests and diseases of citrus were selected as treatments. Assays were conducted to study adult (the most susceptible stage of the parasitoid) exposure to the products; direct contact with immature forms (egg-larva, prepupa, and pupa) inside eggs of the natural host G. aurantianum and of the factitious host, Anagasta kuehniella; and acceptance, by females of the parasitoid, of eggs treated with the products. The products were then classified with regard to the reduction caused in parasitism or emergence percentage, into: 1, harmless (<30%); 2, slightly harmful (30 to 79%); 3, moderately harmful (80 to 99%); and 4, harmful (>99%). The insecticides, miticides, and fungicides tested affected T. atopovirilia parasitism. No differences were observed in T. atopovirilia behavior reared either on the natural or on the factitious host. The least selective products to parasitoid adults were deltamethrin, abamectin, mineral oil, and imidacloprid. As to the immature stages of the parasitoid, all products were harmless, except for mineral oil, which caused a 100% reduction in emergence percentage; abamectin, triflumuron, and imidacloprid were slightly harmful to the egg-larval period. Abamectin and triflumuron had the greatest effect on the parasitism capacity of individuals emerged when the treatment was applied during the parasitoid\'s pupal stage. The products Bacillus thuringiensis, difenoconazole, copper oxychloride, pyraclostrobin, dicofol, spirodiclofen, considered harmless or slightly harmful in all steps of the laboratory assays, can be used in Integrated Pest Management programs, in association with biological control, without being harmful to the parasitoid, and can contribute toward citriculture sustainability.

As plantas reconhecem e produzem substâncias voláteis específicas para a atração de parasitóides após o ataque de um herbívoro, num processo conhecido como defesa indireta. Contudo, a capacidade dessas plantas em processar e liberar novos compostos voláteis sob o ataque de um herbívoro não hospedeiro permanece ainda inexplorado. Em vista disso, o presente trabalho buscou investigar o efeito dos voláteis emitidos por planta de milho sob o ataque de uma praga até então não hospedeira (traça-das-crucíferas, Plutella xylostella) comparado ao daqueles emitidos por planta de milho sob ataque de um hospedeiro tradicional (lagarta-do-cartucho-domilho, Spodoptera frugiperda), sobre esses insetos e seus respectivos parasitóides, Apanteles piceotrichosus e Campoletis flavicincta. Os bioensaios com os parasitóides foram conduzidos em fotofase, sendo oferecidos a estes, voláteis de plantas de milho sadias, voláteis liberados nos intervalos de 0-1, 5-6 e 24-25h após tratamento em fotofase por dano mecânico ou herbivoria simulada, e voláteis liberados no intervalo de 5-6h após esses mesmos tratamentos em escotofase. Já os ensaios com as mariposas foram conduzidos em escotofase, sendo oferecidos a estas, voláteis de plantas de milho sadias, voláteis liberados nos intervalos de 5-6h após tratamento em fotofase por dano mecânico ou herbivoria simulada, e voláteis liberados no intervalo de 0-1 e 24- 25h após esses mesmos tratamentos em escotofase. Adicionalmente, foram testados, para P. xylostella, voláteis de plantas de couve-manteiga sadia, e para A. piceotrichosus, voláteis de couve-manteiga sadia, couve-manteiga danificada mecanicamente e couve-manteiga atacada por lagartas de P. xylostella. As fêmeas de C. flavicincta apresentaram atratividade para voláteis emitidos pelas plantas de milho no intervalo de 5 a 6 horas após tratamento com regurgito de S. frugiperda em escotofase. Curiosamente, não apresentou atratividade pelos voláteis liberados nesse mesmo intervalo de tempo após indução com regurgito em fotofase. As fêmeas acasaladas de S. frugiperda foram atraídas por voláteis de plantas de milho sadia e voláteis liberados nos intervalos de 5-6 e 24-25h após dano mecânico ou tratamento da planta com regurgito deste herbívoro. Porém, preferiu voláteis de plantas sadias aqueles de plantas tratadas com regurgito em fotofase. Esses resultados demonstraram que esses insetos são capazes de discriminar entre misturas de voláteis presentes em seu habitat natural, onde ocorre a relação tritrófica milho (planta hospedeira) S. frugiperda (herbívoro) C. flavicincta (parasitóide). Da mesma forma, na relação couve-manteiga (planta hospedeira) P. xylostella (herbívoro) A. piceotrichosus (parasitóide), as fêmeas de P. xylostella foram atraídas pelos voláteis de couve-manteiga sadia, assim como as fêmeas de A. piceotrichosus foram atraídas pelos voláteis de couve-manteiga atacadas por lagartas deste herbívoro. No caso da simulação de uma nova relação, milho (planta não-hospedeira) P. xylostella (herbívoro) C. flavicincta (parasitóide), tanto fêmeas de A. piceotrichosus, quanto de P. xylostella, não foram capazes de responder aos voláteis de plantas de milho sadias, danificadas mecanicamente e danificadas mecanicamente + regurgito de P. xylostella. A determinação desses mecanismos poderá ser útil para maior compreensão do contexto evolutivo entre plantas e insetos e obtenção de novos avanços no manejo e controle biológico de pragas. / Plants recognize and produce specific volatile substances that attract parasitoids after the herbivore attack, characterizing a process known as indirect defense. However, the ability of these plants in processing and releasing novel volatile compounds elicited by a non-host herbivore attack has been poorly explored. Regarding this, the current study aimed to investigate the effect of volatiles emitted by maize plants under the attack of a pest which is not a host so far, diamondback moth, Plutella xylostella compared to those emitted by a common host, fall armyworm, Spodoptera frugiperda on the behavior response of these insects and their respective parasitoids Apanteles piceotrichosus e Campoletis flavicincta. The bioassays with the parasitoids were conducted during photophase and they were exposed to volatiles from undamaged maize, volatiles released at the time intervals 0-1, 5-6 and 24-25h after the treatment of mechanical damage or simulated herbivory during photophase, and volatiles released at the time interval 5-6h after these same treatments in scotophase. The bioassays with the moths were carried out in scotophase and they were exposed to volatiles from undamaged maize, volatiles released at the time interval 5-6h after the treatment of mechanical damage or simulated herbivory during photophase, and volatiles released at time intervals 0-1 and 24-25h after these same treatments in scotophase. Additionally, for P. xylostella volatiles from undamaged kale were tested while for A. piceotrichosus it was tested volatiles from undamaged kale, mechanically damaged and P. xylostella caterpillar damaged kale. C. flavicincta females were attracted to volatiles emitted by the maize plants at the interval 5-6h after the treatment with the S. frugiperda regurgitate in scotophase. Curiously, they were not attracted to volatiles released at the same time interval after the induction elicited by the regurgitate in the photophase. S. frugiperda mated females were attracted by volatiles from undamaged plants and volatiles released at time intervals 5-6 and 24- 25h after the mechanical damage or treated with the regurgitate of this herbivore. Nevertheless, they preferred the volatiles from undamaged maize to the plants treated with the regurgitate during photophase. These results demonstrated that these insects are able to distinguish among the volatile blends present in their natural habitat where it occurs the tritrophic relationship maize (host plant) S. frugiperda (herbivore) C. flavicincta (parasitoid). In the same way, in the relationship kale (host plant)- P. xylostella (herbivore) A. piceotrichosus (parasitoid), P. xylostella females were attracted by the volatiles of undamaged kale as well as the A. piceotrichosus females were attracted to volatiles emited by caterpillar-damaged kale. In the case of simulating a new relationship, maize (non-host plant) P. xylostella (herbivore) C. flavicincta (parasitoid), A. piceotrichosus females and P. xylostella were not able to respond to undamaged maize, mechanically damaged and mechanically damaged+ P. xylostella regurgitate. The determination of these mechanisms can be useful for a better understanding of the evolution context between plants and insects and for obtaining new advances in the management and biological pest control.

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.

<p>Landscape effects on species performance currently receives much attention. Habitat loss and fragmentation are considered major threats to species diversity. Deciduous forests in southern Sweden are previous wooded pastures that have become species-rich communities appearing as islands in agricultural landscapes, varying in species composition. <i>Actaea spicata</i> is a long-lived plant occurring in these forests. In 150 populations in a 10-km² area, I studied pre-dispersal seed predation, seed dispersal and pollination. I investigated spatio-temporal dynamics of a tritrophic system including<i> Actaea</i>, a specialist seed predator, <i>Eupithecia immundata</i>, and its parasitoids. In addition, effects of biotic context on rodent fruit dispersal and effects of flowering time and flower number on seed set, seed predation and parasitization were studied. Insect incidences of both trophic levels were related to resource population size and small <i>Eupithecia</i> populations were maintained by the rescue effect. There was a unimodal relationship between seed predation and plant population size. Seed predator populations frequently went extinct in small plant populations, resulting in low average seed predation. Parasitoids were present in large plant populations but did not affect seed predator density. Seed predators aggregated at edges, relaxing seed predation in patch interiors. Flowering phenology was unrelated to seed set and insect incidence. A higher flower number did not influence seed predation but was associated with higher seed set and a tendency for a higher parasitization rate. In the study on fruit dispersal more fruits were removed inside than outside populations. Within plant populations more fruits were removed from large aggregations. Overall, this thesis underlines the importance of plant-animal interactions during different phases of the life cycle. The spatial configuration of host plants plays an important role for the outcome of plant-animal interactions and trophic cascades.</p>

Landscape effects on species performance currently receives much attention. Habitat loss and fragmentation are considered major threats to species diversity. Deciduous forests in southern Sweden are previous wooded pastures that have become species-rich communities appearing as islands in agricultural landscapes, varying in species composition. Actaea spicata is a long-lived plant occurring in these forests. In 150 populations in a 10-km2 area, I studied pre-dispersal seed predation, seed dispersal and pollination. I investigated spatio-temporal dynamics of a tritrophic system including Actaea, a specialist seed predator, Eupithecia immundata, and its parasitoids. In addition, effects of biotic context on rodent fruit dispersal and effects of flowering time and flower number on seed set, seed predation and parasitization were studied. Insect incidences of both trophic levels were related to resource population size and small Eupithecia populations were maintained by the rescue effect. There was a unimodal relationship between seed predation and plant population size. Seed predator populations frequently went extinct in small plant populations, resulting in low average seed predation. Parasitoids were present in large plant populations but did not affect seed predator density. Seed predators aggregated at edges, relaxing seed predation in patch interiors. Flowering phenology was unrelated to seed set and insect incidence. A higher flower number did not influence seed predation but was associated with higher seed set and a tendency for a higher parasitization rate. In the study on fruit dispersal more fruits were removed inside than outside populations. Within plant populations more fruits were removed from large aggregations. Overall, this thesis underlines the importance of plant-animal interactions during different phases of the life cycle. The spatial configuration of host plants plays an important role for the outcome of plant-animal interactions and trophic cascades.

One of the large lineages of parasitic wasps, the Cynipoidea, exhibits three distinctly different life modes. Slightly more than half of the about 3000 species are parasitoids in insect larvae, whereas the remaining species are associated with plants, either as gall inducers or as inquilines (guests feeding on plant tissue in galls). The main focus of this thesis has been to identify morphological changes associated with the shifts between life modes. Particular attention was paid to structures believed to be important in gall initiation. Comparative anatomical studies of the egg, larva and venom apparatus were performed, including representatives of parasitoids, gall inducers and inquilines. Examination of gross morphology and ultrastructure revealed that the eggs of the gall inducers are larger and surrounded by a thicker shell than the parasitoid eggs. These differences may be related to the fact that the gall inducer egg contains sufficient egg yolk for the embryo during the entire egg period, whereas the parasitoid egg often absorbs nutrients through the eggshell. Furthermore, the gall inducer egg is probably more exposed to desiccation and therefore a thicker and more resistant eggshell is crucial. Comparing the terminal-instar larvae of about 30 species of parasitoids, gall inducers and inquilines, extensive morphological variation was found, particularly in the head and mouthpart features. The variation was summarized in 33 morphological and one life-history character and parsimony analyses were performed. The resulting phylogenetic estimates were largely in accordance with previous analyses of adult morphology and molecular data. The larval data point to a single origin of the inquilines, in agreement with adult morphology but in conflict with molecular data. The venom apparatus was found to be quite uniform in structure among a sample of 25 species of cynipoid species. It consists of a very short venom duct, a reservoir and a single unbranched venom gland. With few exceptions, the venom apparatus is conspicuously larger relative to the female metasoma in the gall inhabiting species than in the parasitoids. We found little evidence of anatomical structures that could facilitate chemical communication between the gall-inducer embryo and the surrounding plant tissue through the thick eggshell. On the other hand, the enormous venom glands and reservoirs, which are apparently not used for defence, suggest that the adult female plays a significant role in gall induction by injecting secretions into the host plant when laying eggs.

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.

Indirect ecological effects such as associational resistance and resource competition have the potential to affect ecological interactions and influence the structure of ecological communities. Although resource competition is commonly studied, the effects of associational resistance are not as evident if studies are not designed to detect them. Additionally, the relative strengths of different ecological mechanisms ought to be measured in studies, rather than the strength of singular mechanisms. This permits proper attribution of causes and effects in community structure and detection of higher order interactions in a way that naïve reductionism will not. In a series of experiments, I looked at the effects of large-scale addition and removal of Borrichia frutescens on associational resistance of Iva frutescens to the gallformer Asphondylia borrichiae in order to test the mechanism and strength of associational resistance in the system. Additionally, I measured the effects of relative host abundance and interpatch distance of hosts on associational resistance. Finally, I looked at the effect of the presence of stemborers competing with the gall former for host plant resources on parasitism rate and parasitoid guild composition. I found evidence for a strong effect of associational resistance by natural enemies over short distances, although the phenomenon is likely of minor importance in comparison to other factors known to affect gall former population density, such as environmental effects and host plant genotype. Competitors also had a weak effect, reducing mean gall diameter, but not significantly altering total parasitism rate. However, the presence of stemborer competitors did slightly alter the composition of the parasitoid guild. In sum, both associational resistance and competition from stemborers have detectable effects on A. borrichiae, albeit weak ones. Environmental factors, such as soil nitrogen content, are likely much stronger determinants of gall former population size.