Este estudo investigou a ecologia de vespas parasitóides ocorrentes em galhadores Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) em folhas de Lantana camara L. (Verbenaceae). Dessa forma, o primeiro artigo trata da ecologia comportamental da vespa parasitóide Torymus sp. (Torymidae, Hymenoptera), onde foi avaliado como as fêmeas dessa espécie defendem seu recurso de oviposição no hospedeiro. Foi analisado mudanças no comportamento devido à presença de outra fêmea coespecífica no local, ser residente do recurso, tamanho das vespas e número e tamanho das galhas através de filmagens dos experimentos realizados em laboratório. Com esse trabalho observou-se que fêmeas mudam seu comportamento quando estão na presença de um competidor em um território com hospedeiros. A estratégia de ataque foi através da ameaça, na qual suas antenas e asas são levantadas para expulsar o competidor do local. A probabilidade de haver ataques a fêmeas coespecíficas depende do tempo prévio de exploração da galha e da permanência na folha. O interesse no hospedeiro pelas invasoras foi a principal causa de conflitos com a residente. O segundo artigo trata da diversidade de vespas parasitóides e de um ciclo parasita-hospedeiro observado no período de um ano no sistema de L. camara. Foram encontradas nove espécies de vespas parasitóides, divididas em quatro famílias. Ocorreu um ciclo no parasitismo das espécies de vespas sobre seu hospedeiro no ano amostrado, podendo ser observado que são mais elevadas nos meses de Julho a Janeiro e a sobrevivência do hospedeiro foi maior nos meses de Fevereiro a Maio. As estratégias comportamentais de fêmeas de uma vespa parasitóide em relação a seus hospedeiros foram analisadas, bem como um sistema composto de diversas espécies parasitóides e sua variação no tempo. Estas observações podem contribuir para um melhor entendimento da ecologia comportamental e do padrão temporal das vespas parasitoides, e também para futuros programas de controle biológico mais eficientes. / We investigated the ecology of parasitoid wasps attacking Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) galls on leaves of Lantana camara L. (Verbenaceae). The first article discusses the behavioral ecology of the parasitoid wasp Torymus sp. (Torymidae, Hymenoptera), reporting how their females defend oviposition resources on the hosts, changing their behavior due to the presence of a conspecific female in the patch. The identity of the wasp (resident or intruder) on the resource, female size and number and size of galls in the patch were factors studied through analysis of the behaviors revealed by video recordings of the laboratory experiments. Females change their behavior in the presence of a competitor in a territory with hosts. The strategy was to threat, raising their antennae and wings to expel the competitor of the patch. The probability of an attack on a conspecific females depended on the host exploitation time and time spent on the galled leaf. Interest in host by intruders was the main cause for conflicts. The second article reports the diversity of parasitoid wasps and a host-parasite cycle during one year period in the L. camara leaf galls system. Nine parasitoid wasp species were found, divided in four families. A cycle between parasitism and host survival was found during the sampling period. Wasp species are more abundant from July to January and host survival higher from February to May. Behavioral strategies of parasitoid wasp females toward their hosts and conspecifics have been elucidated, with the system composed of different parasitoid wasp species apparently going through an annual cycle of parasitism rate. These observations may contribute to a better understanding of parasitoid behavioral ecology and host-parasitoid dynamics, enabling more efficient future biological control programs.

Este estudo investigou a ecologia de vespas parasitóides ocorrentes em galhadores Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) em folhas de Lantana camara L. (Verbenaceae). Dessa forma, o primeiro artigo trata da ecologia comportamental da vespa parasitóide Torymus sp. (Torymidae, Hymenoptera), onde foi avaliado como as fêmeas dessa espécie defendem seu recurso de oviposição no hospedeiro. Foi analisado mudanças no comportamento devido à presença de outra fêmea coespecífica no local, ser residente do recurso, tamanho das vespas e número e tamanho das galhas através de filmagens dos experimentos realizados em laboratório. Com esse trabalho observou-se que fêmeas mudam seu comportamento quando estão na presença de um competidor em um território com hospedeiros. A estratégia de ataque foi através da ameaça, na qual suas antenas e asas são levantadas para expulsar o competidor do local. A probabilidade de haver ataques a fêmeas coespecíficas depende do tempo prévio de exploração da galha e da permanência na folha. O interesse no hospedeiro pelas invasoras foi a principal causa de conflitos com a residente. O segundo artigo trata da diversidade de vespas parasitóides e de um ciclo parasita-hospedeiro observado no período de um ano no sistema de L. camara. Foram encontradas nove espécies de vespas parasitóides, divididas em quatro famílias. Ocorreu um ciclo no parasitismo das espécies de vespas sobre seu hospedeiro no ano amostrado, podendo ser observado que são mais elevadas nos meses de Julho a Janeiro e a sobrevivência do hospedeiro foi maior nos meses de Fevereiro a Maio. As estratégias comportamentais de fêmeas de uma vespa parasitóide em relação a seus hospedeiros foram analisadas, bem como um sistema composto de diversas espécies parasitóides e sua variação no tempo. Estas observações podem contribuir para um melhor entendimento da ecologia comportamental e do padrão temporal das vespas parasitoides, e também para futuros programas de controle biológico mais eficientes. / We investigated the ecology of parasitoid wasps attacking Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) galls on leaves of Lantana camara L. (Verbenaceae). The first article discusses the behavioral ecology of the parasitoid wasp Torymus sp. (Torymidae, Hymenoptera), reporting how their females defend oviposition resources on the hosts, changing their behavior due to the presence of a conspecific female in the patch. The identity of the wasp (resident or intruder) on the resource, female size and number and size of galls in the patch were factors studied through analysis of the behaviors revealed by video recordings of the laboratory experiments. Females change their behavior in the presence of a competitor in a territory with hosts. The strategy was to threat, raising their antennae and wings to expel the competitor of the patch. The probability of an attack on a conspecific females depended on the host exploitation time and time spent on the galled leaf. Interest in host by intruders was the main cause for conflicts. The second article reports the diversity of parasitoid wasps and a host-parasite cycle during one year period in the L. camara leaf galls system. Nine parasitoid wasp species were found, divided in four families. A cycle between parasitism and host survival was found during the sampling period. Wasp species are more abundant from July to January and host survival higher from February to May. Behavioral strategies of parasitoid wasp females toward their hosts and conspecifics have been elucidated, with the system composed of different parasitoid wasp species apparently going through an annual cycle of parasitism rate. These observations may contribute to a better understanding of parasitoid behavioral ecology and host-parasitoid dynamics, enabling more efficient future biological control programs.

Este estudo investigou a ecologia de vespas parasitóides ocorrentes em galhadores Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) em folhas de Lantana camara L. (Verbenaceae). Dessa forma, o primeiro artigo trata da ecologia comportamental da vespa parasitóide Torymus sp. (Torymidae, Hymenoptera), onde foi avaliado como as fêmeas dessa espécie defendem seu recurso de oviposição no hospedeiro. Foi analisado mudanças no comportamento devido à presença de outra fêmea coespecífica no local, ser residente do recurso, tamanho das vespas e número e tamanho das galhas através de filmagens dos experimentos realizados em laboratório. Com esse trabalho observou-se que fêmeas mudam seu comportamento quando estão na presença de um competidor em um território com hospedeiros. A estratégia de ataque foi através da ameaça, na qual suas antenas e asas são levantadas para expulsar o competidor do local. A probabilidade de haver ataques a fêmeas coespecíficas depende do tempo prévio de exploração da galha e da permanência na folha. O interesse no hospedeiro pelas invasoras foi a principal causa de conflitos com a residente. O segundo artigo trata da diversidade de vespas parasitóides e de um ciclo parasita-hospedeiro observado no período de um ano no sistema de L. camara. Foram encontradas nove espécies de vespas parasitóides, divididas em quatro famílias. Ocorreu um ciclo no parasitismo das espécies de vespas sobre seu hospedeiro no ano amostrado, podendo ser observado que são mais elevadas nos meses de Julho a Janeiro e a sobrevivência do hospedeiro foi maior nos meses de Fevereiro a Maio. As estratégias comportamentais de fêmeas de uma vespa parasitóide em relação a seus hospedeiros foram analisadas, bem como um sistema composto de diversas espécies parasitóides e sua variação no tempo. Estas observações podem contribuir para um melhor entendimento da ecologia comportamental e do padrão temporal das vespas parasitoides, e também para futuros programas de controle biológico mais eficientes. / We investigated the ecology of parasitoid wasps attacking Schismatodiplosis lantanae Rübsaamen, 1916 (Cecidomyiidae) galls on leaves of Lantana camara L. (Verbenaceae). The first article discusses the behavioral ecology of the parasitoid wasp Torymus sp. (Torymidae, Hymenoptera), reporting how their females defend oviposition resources on the hosts, changing their behavior due to the presence of a conspecific female in the patch. The identity of the wasp (resident or intruder) on the resource, female size and number and size of galls in the patch were factors studied through analysis of the behaviors revealed by video recordings of the laboratory experiments. Females change their behavior in the presence of a competitor in a territory with hosts. The strategy was to threat, raising their antennae and wings to expel the competitor of the patch. The probability of an attack on a conspecific females depended on the host exploitation time and time spent on the galled leaf. Interest in host by intruders was the main cause for conflicts. The second article reports the diversity of parasitoid wasps and a host-parasite cycle during one year period in the L. camara leaf galls system. Nine parasitoid wasp species were found, divided in four families. A cycle between parasitism and host survival was found during the sampling period. Wasp species are more abundant from July to January and host survival higher from February to May. Behavioral strategies of parasitoid wasp females toward their hosts and conspecifics have been elucidated, with the system composed of different parasitoid wasp species apparently going through an annual cycle of parasitism rate. These observations may contribute to a better understanding of parasitoid behavioral ecology and host-parasitoid dynamics, enabling more efficient future biological control programs.

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.

Ecological network theory has developed from studies of static, binary trophic relationships to the analysis of quantitative, dynamic communities consisting of multiple link-types. Particularly, work has focused on the dynamic nature of ecological networks, which maintains stability in complex communities. However, there are few in situ network-level studies of the determinants of temporal and spatial variation in community structure. This thesis utilises data from a 10-year study of a host-parasitoid network and a collaborative study in an applied ecological setting to identify individual level factors important to network structure. The work aims towards an empirical, predictive framework linking adaptive foraging behaviour to ecological network structure. The results show that condition-dependent foraging behaviours structure host-parasitoid networks. The realised niches of the studied parasitoid species were generally biased towards larger host species and condition-dependent sex ratio allocation increased the likelihood that females would eclose from relatively larger hosts and males from relatively smaller hosts, which resulted in sex ratios deviating from Hamiltonian (50:50) predictions. Further, both of these aspects of behaviour are plastic, where parasitoid behaviour responded to environmental heterogeneity. Particularly, host preference behaviour conformed to an egg-/time-limitation framework, where the size dependency of the behaviour is greater when individuals have a greater likelihood of being egg-limited. Both the size-dependency and the plasticity of these behaviours differed significantly between secondary parasitoid species. This species identity effect interacted with landscape heterogeneity, which may explain some inter- and intra-specific variation in network structure. With respect to applied ecology, the results show that the benefits of natural vegetation for pest control are dependent upon the dispersal capabilities and the diet breadth of the pest and its natural enemies. The findings are evaluated towards a predictive framework for understanding the effects of future climate change on community structure and stability. We consider this framework in terms of applied ecology, particularly pest control ecosystem services provided by natural vegetation in an agricultural environment. The synergistic nature of the multiple determinants of network structure found in this thesis suggest that future studies should focus on the whole network, which is not necessarily the sum of its parts.

In host-parasitoid systems, there is a continuous coevolutionary arms race where each species imposes a strong selection pressure on the other. The host needs to develop defence strategies in order to escape parasitism and the parasitoid must evolve counter-defence strategies in order to overcome the host’s immune defence and successfully reproduce. This makes host-parasitoid systems excellent model systems for understanding evolutionary processes underlying host race formation and speciation. In order to gain a better understanding of the complexity of host-parasitoid interactions several aspects must be considered, such as search behaviour and host selection in the parasitoid, the development of immune responses in the host and counter-defence strategies in the parasitoid. In this thesis, I investigate interactions and coevolution in a natural host-parasitoid system, consisting of five species of Galerucella leaf beetles and three species of Asecodes parasitoids, by combining behavioural ecology with chemical ecology and immunology. In the studies performed, I found that pheromone production and responses in the beetles are connected to the phylogenetic relatedness between the Galerucella species (Paper I). I found no evidence that Asecodes exploits the adult pheromone to locate host larvae, but observed an ability in the parasitoids to distinguish a better host from a less suitable one based on larval odors (Paper II). The studies also revealed large differences in immune competence between the Galerucella species, which were linked to differences in hemocyte composition in the beetle larvae (Paper III, IV). Further, the results suggest that parasitism success in polyphagous Asecodes is strongly affected by former host species of the parasitoid (Paper IV). In conclusion, the results of this thesis suggest an on-going evolution in both parasitoid virulence and host immune responses in the Asecodes-Galerucella system. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 4: Manuscript.</p>

L'espèce Plutella xylostella (L.) (Lepidoptera : Plutellidae) défoliatrice des choux constitue surtout un problème dans les régions tropicales et subtropicales. La lutte chimique a rapidement montré ses limites du fait de l'apparition de résistance dans les populations. Des moyens de lutte alternatifs ont été mis en place, impliquant principalement des insectes parasitoïdes, parmi lesquels Oomyzus sokolowskii (Kurdjumov) (Hymenoptera : Eulophidae) et Cotesia vestalis (Haliday) (Hymenoptera : Braconidae) qui sont les plus couramment utilisés en raison de leur spécificité envers P. xylostella. Afin de contribuer à une meilleure connaissance du contrôle de la teigne en région tropicale, nous avons étudié les relations hôteparasitoïde entre P. xylostella et O. sokolowskii d’une part, et P. xylostella et C. vestalis d’autre part, en conditions de laboratoire et sur le terrain au Sénégal et au Bénin. Au Sénégal, quatre espèces de parasitoïdes sont présentes sur les chenilles : O. sokolowskii, Apanteles litae, C. vestalis et Brachymeria citrae. Au Bénin, seule l'espèce C. vestalis est présente. Au Sénégal comme au Bénin, les facteurs climatiques contribuent au développement de la teigne et les précipitations ne régulent pas les populations du ravageur. Dans ces deux pays, la teigne n’est pas contrôlée par ses ennemis naturels. La lutte biologique par conservation y est à prendre en considération et l’utilisation de plantes compagnes cultivées en association avec le chou peut être envisagée pour réduire les populations de la teigne. Les études en laboratoire ont montré qu’Oomyzus sokolowskii est un parasitoïde larvo-nymphal performant. ConcernantC. vestalis, les femelles détectent et reconnaissent leur hôte grâce aux lipides cuticulaires émis par les chenilles. Des marqueurs moléculaires (isozymes et ISSR) ont confirmé une forte variabilité entre les populations de P. xylostella à l’échelle mondiale, les populations d’Australie et du Japon étant très différentes des autres et formant deux groupes distincts. La structuration des populations semble influencée par le type de climat : tropical et non tropical. / The diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera: Plutellidae), is the most destructive pest of Brassicaceae worldwide and poses particularly acute problems in tropical areas. Chemical control is impaired by multiple-insecticide resistance in this species. Alternative methods are based on biological control by parasitoids, such as Oomyzussokolowskii (Kurdjumov) (Hymenoptera: Eulophidae) and Cotesia vestalis (Haliday) (Hymenoptera: Braconidae), which are commonly used due to their specificity towards DBM. To help to improve the biocontrol of the moth in the tropics, we studied host-parasitoid relationships between P. xylostella and these two parasitoids under both laboratory and field conditions in Senegal and Benin. In both countries, climatic conditions are favourable for the development of DBM and rainfall does not limit populations of this pest. In Senegal, four parasitoid species are present on DBM larvae: O. sokolowskii, C. vestalis, Apanteles litae, and Brachymeria citrae. In Benin, C. vestalis is largely dominant. In neither of these countries, the moth is sufficiently controlled by natural enemies. Conservation biological control might be combined with the use of companion plants in cabbage crops to reduce DBM populations. Laboratory studies have shown that O. sokolowskii is an efficient larval-pupal parasitoid. In C. vestalis, females detect and recognize their host using cuticular lipids produced by the caterpillar. Studies of molecular markers (isozymes and ISSR) have confirmed high variability among DBM populations around the world, those from Australia and Japan being distinct and very different from any other population. Population structure seems to be influenced by the type of climate (tropical vs. non-tropical).

La manipulation du comportement de l'hôte est une stratégie couramment utilisée chez les organismes parasites, et ce afin de favoriser leur transmission et/ou leur survie. La compréhension d'une telle stratégie est au cœur de plusieurs disciplines appliquées de la parasitologie, telles que l'épidémiologie et la médecine, mais est aussi d'une grande importance d'un point de vue écologique et évolutif. Dans le cadre de cette thèse, notre objectif était d'améliorer l'état des connaissances sur un type de manipulation encore très peu connu : la manipulation garde du corps, une stratégie initialement décrite chez les insectes parasitoïdes. À la suite de leur développement larvaire, certaines guêpes parasitoïdes (Braconidae) ont la capacité d'usurper le comportement de leur hôte et l'obligent ainsi à se comporter comme un véritable garde du corps pour protéger leur cocon des ennemis naturels. Basé sur le système associant la guêpe parasitoïde Dinocampus coccinellae et un de ses hôtes la coccinelle maculée Coleomegilla maculata, le présent travail aborde cinq aspects particuliers de la manipulation parasitaire : i) Efficacité de la protection par un garde du corps et coûts liés à une telle stratégie, ii) Contraintes énergétiques pour les parasites manipulateurs, iii) Diversité et l'évolution de la manipulation garde du corps, iv) Survie des hôtes à la manipulation par un parasitoïde et v) Effets de la nutrition de l'hôte sur une association hôte–parasitoïde manipulateur. Nos résultats apportent plusieurs éléments de réponse quant à l'écologie et l'évolution de la manipulation garde du corps, et notamment en ce qui a trait aux coûts énergétiques associés à une telle stratégie ainsi qu'au rôle de la ressource dans cette étroite association. De nouvelles perspectives de recherche y sont proposées afin d'encourager d'autres travaux dans cette voie. / Host behavioral manipulation, a widespread strategy among parasitic organisms, has evolved to favor the transmission and/or survival of parasites. Understanding such a strategy is at the heart of applied aspects of parasitology, such as epidemiology and medicine, but is also important for evolutionary and ecological reasons. The aim of this study was to improve our knowledge on a category of manipulation that remains poorly documented: the bodyguard manipulation, initially described on parasitoid insects. Following their larval development, certain parasitic wasps (Braconidae) are able to usurp the behavior of their host and force them to serve as bodyguards to protect parasitoid cocoon against natural enemies. Studying the association between the parasitic wasp Dinocampus coccinellae and one of its host the spotted lady beetle Coleomegilla maculata, this work is structured in five axes: i) Efficiency of the bodyguard protection and associated costs, ii) Energetic constraints for manipulative parasites, iii) Diversity and evolution of bodyguard manipulation, iv) Host survival from parasitism and v) Host nutrition effects on a host–manipulative parasitoid association. Our results provide original findings about the ecology and evolution of the bodyguard manipulation, for instance the energetic costs associated with host manipulation or the role of host nutrition in this type of association. New perspectives are identified in order to stimulate future research in this area.

This dissertation is concerned with effects dispersal and forest structure on forest insect population dynamics, and with identifying generating processes by comparing observed patterns to model predictions. In chapter 2, we investigated effects of changing forest landscape patterns on integro-difference models of host-parasitoid population dynamics. We demonstrated that removing habitat can increase herbivore density when herbivores don't disperse far, and parasitoids disperse further, due to differences in dispersal success between trophic levels. This is a novel potential explanation for why forest fragmentation increases the duration of forest tent caterpillar outbreaks. To better understand spatial model behaviour, we proposed a new local variation of the dispersal success approximation. The approximation successfully predicts effects of habitat loss and fragmentation on realistically complex landscapes, except when outbreak cycle amplitude is very large. Local dispersal success is useful in part because parameters can be estimated from widely available habitat data. In chapter 3, we investigated how well a discretized integro-difference model of mountain pine beetle population dynamics predicted the occurrence of new infestations in British Columbia. We found that a model with a large dispersal kernel, and high emigration from new, low severity infestations yielded the best predictions. However, we do not believe this to be convincing evidence that many beetles disperse from new, low severity infestations. Rather, we argued that differences in habitat quality, detection errors, and Moran effects can all confound dispersal patterns, making it difficult to infer dispersal parameters from observed infestation patterns. Nonetheless, predicting infestation risk is useful, and large kernels improve predictions. In chapter 4, we used generalized linear mixed models to characterize spatial and temporal variation in the propensity of jack pine trees to produce pollen cones, and account for confounding effects on the relationship between pollen cone production and previous defoliation by jack pine budworm. We found effects of stand age, and synchronous variation in pollen cone production among years. Accounting for background patterns in pollen cone production clarified that pollen cone production declines in with previous defoliation, as expected.

This dissertation is concerned with effects dispersal and forest structure on forest insect population dynamics, and with identifying generating processes by comparing observed patterns to model predictions. In chapter 2, we investigated effects of changing forest landscape patterns on integro-difference models of host-parasitoid population dynamics. We demonstrated that removing habitat can increase herbivore density when herbivores don't disperse far, and parasitoids disperse further, due to differences in dispersal success between trophic levels. This is a novel potential explanation for why forest fragmentation increases the duration of forest tent caterpillar outbreaks. To better understand spatial model behaviour, we proposed a new local variation of the dispersal success approximation. The approximation successfully predicts effects of habitat loss and fragmentation on realistically complex landscapes, except when outbreak cycle amplitude is very large. Local dispersal success is useful in part because parameters can be estimated from widely available habitat data. In chapter 3, we investigated how well a discretized integro-difference model of mountain pine beetle population dynamics predicted the occurrence of new infestations in British Columbia. We found that a model with a large dispersal kernel, and high emigration from new, low severity infestations yielded the best predictions. However, we do not believe this to be convincing evidence that many beetles disperse from new, low severity infestations. Rather, we argued that differences in habitat quality, detection errors, and Moran effects can all confound dispersal patterns, making it difficult to infer dispersal parameters from observed infestation patterns. Nonetheless, predicting infestation risk is useful, and large kernels improve predictions. In chapter 4, we used generalized linear mixed models to characterize spatial and temporal variation in the propensity of jack pine trees to produce pollen cones, and account for confounding effects on the relationship between pollen cone production and previous defoliation by jack pine budworm. We found effects of stand age, and synchronous variation in pollen cone production among years. Accounting for background patterns in pollen cone production clarified that pollen cone production declines in with previous defoliation, as expected.