There is anecdotal evidence that populations of the New Zealand endemic red admiral butterfly Bassaris gonerilla (F.) have declined since the early 1900s. This decline has been associated with the introduction of the generalist pupal parasitoids Pteromalus puparum (L.) and Echthromorpha intricatoria (F.). The former was deliberately introduced for the biological control of the cabbage white butterfly (Pieris rapae (L.)); the latter is an adventitious arrival from Australia. The objective of this thesis was to quantify, using population models, the effect that P. puparum is having on B. gonerilla abundance. Population monitoring and a phenology model (based on temperature-related development rates) indicated that B. gonerilla has two full generations and one partial generation per summer in the Banks Peninsula region of New Zealand. B. gonerilla abundance was greatly reduced in drought summers, which was probably due to the negative effects of drought on the quality and quantity of the larval host plant Urtica ferox Forst. A life table study showed that egg parasitism by the unidentified scelionid Telenomus sp. was the largest mortality factor for the pre-imaginal stages of B. gonerilla, followed by "disappearance" mortality (predation and dispersal) in the larval stages. Pupal mortality due to P. puparum was lower compared with that caused by E. intricatoria, with 1-19% and 20-30% of pupae being parasitised by P. puparum and E. intricatoria, respectively. Collection of B. gonerilla pupae from the Christchurch, Dunedin and Wellington areas confirmed higher rates of percentage parasitism by E. intricatoria. B. gonerilla collected from the Banks Peninsula had a 50: 50 sex ratio and lifetime fecundity was estimated in the laboratory as 312 eggs per female. There was no evidence of density-dependent parasitism of B. gonerilla pupae by P. puparum in the field, although there was a significant positive relationship between life table estimates of E. intricatoria parasitism and B. gonerilla pupal abundance. Larval dispersal from the host plant showed a positive relationship with larval instar but no relationship with larval density. Rates of change in B. gonerilla adult abundance between generations within a year showed evidence of density dependence, and this negative feedback was stronger in a drought year. A discrete-time model for B. gonerilla population dynamics was constructed which had two summer generations per year and a partial overwintering generation. The model showed that the presence of this overwintering generation provides a temporal refuge from high levels of E. intricatoria parasitism. Removal of parasitoid mortality from the model suggested that P. puparum was suppressing B. Gonerilla populations on the Banks Peninsula by 5% and E. intricatoria by 30%. An important assumption of the model was that parasitism rates were independent of B. gonerilla density. This assumption appears valid for P. puparum parasitism, but may not be valid for E. intricatoria; therefore the estimated suppression levels due to this adventive parasitoid should be viewed with some caution. It is too soon to generalise on what determines the magnitude of non-target effects by arthropod biocontrol agents, this being only the second study to quantify effects at a population level. However, in this case retrospective analysis has shown that the impact of non-target parasitism by P. puparum on B. gonerilla abundance has been small. There is anecdotal evidence that populations of the New Zealand endemic red admiral butterfly Bassaris gonerilla (F.) have declined since the early 1900s. This decline has been associated with the introduction of the generalist pupal parasitoids Pteromalus puparum (L.) and Echthromorpha intricatoria (F.). The former was deliberately introduced for the biological control of the cabbage white butterfly (Pieris rapae (L.)); the latter is an adventitious arrival from Australia. The objective of this thesis was to quantify, using population models, the effect that P. puparum is having on B. gonerilla abundance. Population monitoring and a phenology model (based on temperature-related development rates) indicated that B. gonerilla has two full generations and one partial generation per summer in the Banks Peninsula region of New Zealand. B. gonerilla abundance was greatly reduced in drought summers, which was probably due to the negative effects of drought on the quality and quantity of the larval host plant Urtica ferox Forst.. A life table study showed that egg parasitism by the unidentified scelionid Telenomus sp. was the largest mortality factor for the pre-imaginal stages of B. gonerilla, followed by "disappearance" mortality (predation and dispersal) in the larval stages. Pupal mortality due to P. puparum was lower compared with that caused by E. intricatoria, with 1-19% and 20-30% of pupae being parasitised by P. puparum and E. intricatoria, respectively. Collection of B. gonerilla pupae from the Christchurch, Dunedin and Wellington areas confirmed higher rates of percentage parasitism by E. intricatoria. B. gonerilla collected from the Banks Peninsula had a 50: 50 sex ratio and lifetime fecundity was estimated in the laboratory as 312 eggs per female. There was no evidence of density-dependent parasitism of B. gonerilla pupae by P. puparum in the field, although there was a significant positive relationship between life table estimates of E. intricatoria parasitism and B. gonerilla pupal abundance. Larval dispersal from the host plant showed a positive relationship with larval instar but no relationship with larval density. Rates of change in B. gonerilla adult abundance between generations within a year showed evidence of density dependence, and this negative feedback was stronger in a drought year. A discrete-time model for B. gonerilla population dynamics was constructed which had two summer generations per year and a partial overwintering generation. The model showed that the presence of this overwintering generation provides a temporal refuge from high levels of E. intricatoria parasitism. Removal of parasitoid mortality from the model suggested that P. puparum was suppressing B. Gonerilla populations on the Banks Peninsula by 5% and E. intricatoria by 30%. An important assumption of the model was that parasitism rates were independent of B. gonerilla density. This assumption appears valid for P. puparum parasitism, but may not be valid for E. intricatoria; therefore the estimated suppression levels due to this adventive parasitoid should be viewed with some caution. It is too soon to generalise on what determines the magnitude of non-target effects by arthropod biocontrol agents, this being only the second study to quantify effects at a population level. However, in this case retrospective analysis has shown that the impact of non-target parasitism by P. puparum on B. gonerilla abundance has been small.

Cette thèse porte sur l’analyse des dimensions spatiale et temporelle du fonctionnement et de la diversité taxonomique et fonctionnelle d’une guilde. Au sein d’une guilde, les espèces exploitent un même type de ressources; pour cette raison, les niches écologiques fondamentales des membres d’une guilde sont similaires et une relation de compétition interspécifique se met en place si la ressource partagée est limitante. Quatre questions principales sur le fonctionnement d’une guilde ont été abordées : (1) à quelles échelles spatio-temporelles se structurent la guilde et les ressources exploitées, (2) quelles sont les influences respectives des filtres environnementaux et des interactions interspécifiques sur les traits fonctionnels des membres de la guilde, (3) quelle est la stratégie d’exploitation de ressources d’un membre de la guilde face à une faible disponibilité de ressources et (4) quel est l’impact du contexte climatique sur la structure d’une guilde et de son réseau trophique, et sur le degré de spécialisation écologique des espèces de la guilde sur la ressource. Le modèle biologique de cette étude a été une guilde de parasitoïdes de pucerons de céréales (Hymenoptera : Braconidae : Aphidiinae). Ces parasitoïdes s’attaquent aux pucerons (Homoptera : Aphididae) inféodés aux céréales dans les agroécosystèmes. Les variations d’abondance relative des parasitoïdes et de leurs hôtes ont été importantes aux échelles interrégionale et interannuelle, mais très faibles à l’échelle intra-régionale. La divergence des traits fonctionnels des parasitoïdes s’est maintenue sur trois régions et deux années, et le contexte local a influencé les traits de la guilde dans son ensemble. Pendant la saison hivernale, une période marquée par une faible densité de pucerons, le parasitoïde Aphidius rhopalosiphi a présenté des stratégies contrastées pour maximiser sa fitness en exploitant les hôtes Sitobion avenae et Rhopalosiphum padi, mais une forte spécialisation écologique sur le terrain a été observée en présence d’une espèce compétitrice : Aphidius avenae. Cette présence a pu être corrélée à une hausse des températures hivernales. / This thesis is an analysis of the spatial and temporal dimensions of the functioning and taxonomic and functional diversity of a guild. In a guild, species exploit the same type of resources; consequently, fundamental ecological niches of guild members are similar and an interspecific competitive relationship is established if shared resources are limiting. Four main questions on the functioning of a guild are addressed: (1) the spatiotemporal scales to which guilds the guild and its exploited resources are structures, (2) the respective weight of environmental filtering and interspecific interactions on the configuration of realized niches of guild members, (3) the strategy of resource exploitation in a guild member facing low resource availability and (4) the impact of climatic context on the structure of a guild and its food web, and the degree of specialisation on resources. The biological model chosen in this study was a cereal aphid parasitoid guild (Hymenoptera: Braconidae: Aphidiinae). Those parasitoids attack aphids (Homoptera: Aphididae) in agroecosystems, which are anthropised environments marked by high disturbance rates and in which host resources are variable in density and in quality. Relative abundance variations in parasitoids and in their hosts were considerable at interregional and interannual scales, but weak at the intra-regional scale. Functional trait divergence in parasitoids was maintained across three regions and two years, and regional and annual environmental contexts influenced guild traits as a whole. In winter, a period marked by low aphid density, the parasitoid Aphidius rhopalosiphi presented contrasted strategies to maximise fitness whilst exploiting Sitobion avenae and Rhopalosiphum padi hosts, but a high degree of ecological specialisation in the field was observed in the presence of a competitive species, Aphidius avenae. This presence was correlated to an increase in winter temperatures.

Chez la guêpe parasitoïde Venturia canescens, des particules virales dépourvues d'ADN appelées VLP (pour "Virus-like Particules") sont produites spécifiquement dans les ovaires et tapissent le chorion des oeufs qui sont injectés dans la chenille hôte. Les VLP ont une fonction immunosuppressive pour l'hôte parasité et permettent ainsi la survie des oeufs du parasitoïde. Ces VLP résultent de l’intégration d’un nudivirus dans le génome de l’ancêtre de la guêpe, nudivirus qui a été ensuite domestiqué pour former des liposomes viraux capables de véhiculer dans l’hôte des protéines de virulence d'origine cellulaire. L’étude réalisée au cours de cette thèse a eu pour objet, d’une part, d'étudier les mécanismes de domestication virale qui ont conduit au virus symbiotique endogène actuel nommé VcENV (pour V. canescens endogenous nudivirus) et d’autre part, d'apporter des éléments de réponse sur le processus de morphogénèse et le mode d'action parasitaire des VLP. / Viral particles devoid of DNA called VLPs (for Virus-Like Particles) are specifically produced in the ovaries of the parasitoid wasp Venturia canescens and line the chorion of the wasp’s eggs injected into the host caterpillar. VLPs are immunosuppressive and allow parasitoid eggs survival. These VLPs result from the integration of a nudivirus into the wasp ancestor genome, nudivirus which was then domesticated to form viral liposomes capable of carrying, into the host, virulence proteins of cellular origin. The aim of the study carried out during this thesis was, first, to analyze the viral domestication mechanisms that led to the current endogenous symbiotic virus called VcENV (for V. canescens endogenous nudivirus) and secondly to provide some answers on VLPs morphogenesis process and parasitic mode of action.

In this study, buckwheat (Fagopyrum esculentum Moench) and alyssum (Lobularia maritima (L.)) flowers were used to examine the effect of floral resources on the efficacy of the leafroller parasitoid Dolichogenidea tasmanica (Cameron) in vineyards. This was done by assessing the influence of these flowers on parasitoid abundance and parasitism rate, and by investigating the consequences of this for leafroller abundance. In laboratory experiments, alyssum flowers were used to investigate the effect of floral food on the longevity, fecundity and sex ratio of D. tasmanica. Dolichogenidea tasmanica comprised more than 95 % of parasitoids reared from field collected leafrollers in this study. The abundance of D. tasmanica during the 1999-2000 growing season was very low compared with previous studies, possibly due to the very low abundance of its leafroller hosts during the experiment. The number of males of this species on yellow sticky traps was increased (although not significantly) when buckwheat flowers were planted in a Marlborough vineyard; however, the number of female D. tasmanica on traps was no greater with flowers than without. The abundance of another leafroller parasitoid, Glyptapanteles demeter (Wilkinson)(Hymenoptera: Braconidae), on traps was also not significantly affected by the presence of buckwheat flowers, although females of this species were caught in greater numbers in the control than in buckwheat plots. Naturally-occurring leafrollers were collected from three vineyard sites in Marlborough, and one in Canterbury during the 2000-2001 season to assess the effect of buckwheat and alyssum flowers on parasitism rate. Parasitism rate more than doubled in the presence of buckwheat at one of the Marlborough vineyards, but alyssum had no effect on parasitism rate in Canterbury. A leafroller release/recover method, used when naturally-occurring leafrollers were too scarce to collect, was unable to detect any effect of buckwheat or alyssum on parasitism rate. Mean parasitism rates of approximately 20 % were common in Marlborough, although rates ranged from 0 % to 45 % across the three vineyard sites in that region. In Canterbury in April, mean parasitism rates were approximately 40 % (Chapter 4). Rates were higher on upper canopy leaves (40-60 %) compared with lower canopy leaves and bunches (0-25 %). Leafroller abundance was apparently not affected by the presence of buckwheat in Marlborough, or alyssum in Canterbury. Buckwheat did, however, significantly reduce the amount of leafroller evidence (webbed leafroller feeding sites on leaves or in bunches) in Marlborough, suggesting that the presence of these flowers may reduce leafroller populations. Leafrollers infested less than 0.1 % of Cabernet Sauvignon leaves throughout the 1999-2000 growing season, but increased in abundance in bunches to infest a maximum of 0.5 % of bunches in late March in Marlborough. In Pinot Noir vines in the 2000-2001 season, leafroller abundance was also low, although sampling was not conducted late in the season when abundance reaches a peak. In Riesling vines in Canterbury, between 1.5 % and 2.5 % of bunches were infested with leafrollers in April. In the laboratory, alyssum flowers significantly increased the longevity and lifetime fecundity of D. tasmanica compared with a no-flower treatment. However, daily fecundity was not increased by the availability of food, suggesting that the greater lifetime fecundity was related to increases in longevity. Parasitoids were also able to obtain nutrients from whitefly honeydew, which resulted in similar longevity and daily fecundity to those when alyssum flowers were present. The availability of food had a significant effect on the offspring sex ratio of D. tasmanica. Parasitoids reared from naturally-occurring leafrollers produced an equal sex ratio, assumed to be the evolutionarily stable strategy (ESS) for this species. In the laboratory, this ESS was observed only when parasitoids had access to alyssum flowers. Without food, or with honeydew only, sex ratios were strongly male-biased. In the field, floral resources affected the sex ratio of D. tasmanica only when this species was reared from leafrollers released and recovered in Marlborough. In that experiment, buckwheat shifted the sex ratio in favour of female production from the equal sex ratio found in control plots. No firm explanations can be given to account for these results, due to a lack of research in this area. Possible mechanisms for the changes in sex ratio with flowers are discussed. This study demonstrated that flowers are an important source of nutrients for D. tasmanica, influencing the longevity, fecundity and offspring sex ratio of this species. However, only some of the field experiments were able to show any positive effect of the provision of floral resources on parasitoid abundance or parasitism rate. More information is needed on the role these parasitoids, and other natural enemies, play in regulating leafroller populations in New Zealand vineyards, and on how they use floral resources in the field, before recommendations can be made regarding the adoption of this technology by growers.

Arthropod species that have the potential to damage crops are food resources for communities of predators and parasitoids. From an agronomic perspective these species are pests and biocontrol agents respectively, and the relationships between them can be important determinants of crop yield and quality. The impact of biocontrol agents on pest populations may depend on the availability of other food resources in the agroecosystem. A scarcity of such resources may limit biological control and altering agroecosystem management to alleviate this limitation could contribute to pest management. This is a tactic of ‘conservation biological control’ and includes the provision of flowers for species that consume prey as larvae but require floral resources in their adult stage. The use of flowers for pest management requires an understanding of the interactions between the flowers, pests, biocontrol agents and non-target species. Without this, attempts to enhance biological control might be ineffective or detrimental. This thesis develops our understanding in two areas which have received relatively little attention: the role of flowers in biological control by true omnivores, and the implications of flower use by fourth-trophic-level life-history omnivores. The species studied were the lacewing Micromus tasmaniae and its parasitoid Anacharis zealandica. Buckwheat flowers Fagopyrum esculentum provided floral resources and aphids Acyrthosiphon pisum served as prey. Laboratory experiments with M. tasmaniae demonstrated that although prey were required for reproduction, providing flowers increased survival and oviposition when prey abundance was low. Flowers also decreased prey consumption by the adult lacewings. These experiments therefore revealed the potential for flowers to either enhance or disrupt biological control by M. tasmaniae. Adult M. tasmaniae were collected from a crop containing a strip of flowers. Analyses to determine the presence of prey and pollen in their digestive tracts suggested that predation was more frequent than foraging in flowers. It was concluded that the flower strip probably did not affect biological control by lacewings in that field, but flowers could be significant in other situations. The lifetime fecundity of A. zealandica was greatly increased by the presence of flowers in the laboratory. Providing flowers therefore has the potential to increase parasitism of M. tasmaniae and so disrupt biological control. A. zealandica was also studied in a crop containing a flower strip. Rubidium-marking was used to investigate nectar-feeding and dispersal from the flowers. In addition, the parasitoids’ sugar compositions were determined by HPLC and used to infer feeding histories. Although further work is required to develop the use of these techniques in this system, the results suggested that A. zealandica did not exploit the flower strip. The sugar profiles suggested that honeydew had been consumed by many of the parasitoids. A simulation model was developed to explore the dynamics of aphid, lacewing and parasitoid populations with and without flowers. This suggested that if M. tasmaniae and A. zealandica responded to flowers as in the laboratory, flowers would only have a small effect on biological control within a single period of a lucerne cutting cycle. When parasitoids were present, the direct beneficial effect of flowers on the lacewing population was outweighed by increased parasitism, reducing the potential for biological control in future crops. The results presented in this thesis exemplify the complex interactions that may occur as a consequence of providing floral resources in agroecosystems and re-affirm the need for agroecology to inform the development of sustainable pest management techniques.

Landscape heterogeneity is thought to differ among farm management types (i.e. organic and conventional), and this difference is hypothesized to result in variations in pest control by natural enemies. However, it is unclear if these variations in pest control are driven by landscape structure or by farm management practices themselves. Remotely sensed datasets were used to describe the landscape structure surrounding a group of organic and conventional caneberry farms in Oregon and Washington that have different leafroller parasitism rates attributed to farm management type. A finer scale survey was done at one of the farms using the remotely sensed data as well as field surveys. Landscape metrics of diversity, richness and percent non-crop were used to describe the landscapes surrounding the farm fields at scales ranging from 0.05 km to 5.00 km for the large scale study, and 0.05 km to 0.20 km for the fine scale study. In the fine scale study, data on parasitoid species assemblages, diversity, and parasitism rate were collected and analyzed against the calculated landscape metrics spatially and seasonally. The purpose of this study was to quantify effects of farm management type on habitat structure, effect of habitat structure on leafroller parasitism rate, and to access correlations between landscape metrics calculated at the landscape and field scale. Overall, the farms were embedded in a landscape that was broadly similar, with very few differences in landscape structure occurring between organic and conventional farms. Organic farms had higher vegetation height class diversity at the largest scale compared to conventional farms, while conventional farms had significantly higher percent non-crop area compared to organic farms. There was no significant effect of any of the calculated landscape metrics on parasitism rates. In the field scale study, no correlations were found between habitat metrics and parasitism rates, or between field based metrics and those calculated at the landscape scale. The results of this study suggest that conventional and organic caneberry farms in the Willamette Valley are broadly similar in the habitat conditions they provide parasitoids. This suggests that management changes to pesticide use alone could increase levels of leafroller biological control on conventional farms to levels that are comparable to those seen on organic farms. Our comparisons of the landscape scale and field scale landscape metrics showed no connection, this suggests that direct comparisons cannot be made with these particular metrics at these very different scales. Rather than comparing these types of data, it may be more useful to combine them in order to increase the resolution and predictive power of remotely sensed data for describing landscapes at broad scales. / Graduation date: 2013

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