This study investigated understorey management in Canterbury, New Zealand, apple orchards for the enhancement of populations of Dolichogenidea tasmanica (Cameron) (Braconidae) for leafroller (Lepidoptera: Tortricidae) biological control. The first objective was to determine the influence of understorey plants on the abundance of D. tasmanica and leafroller parasitism, and to investigate the mechanisms behind this influence. The second was to determine the most suitable understorey plants in terms of their ability to enhance parasitoid abundance, leafroller parasitism, parasitoid longevity, parasitoid fecundity and its ability to not benefit leafroller. Results from three consecutive field trials showed that buckwheat (Fagopyrum esculentum Moench), coriander (Coriandrum sativum L.), alyssum (Lobularia maritima (L.) Desv), and, to a lesser extent, broad bean (Vicia faba L.), enhanced parasitoid abundance and leafroller parasitism. The mechanisms behind the effects of understorey plants had previously been unexplored. However, results here showed that it was the flowers or the buckwheat that 'attracted' the parasitoid to the plant and not the shelter, aphids or microclimate that the plant may also provide. Providing flowering plants in the orchard understorey also increased immigration of parasitoids and enhanced parasitoids and enhanced parasitoid longevity and fecundity in the laboratory. In contrast, the understorey plants had no influence on the female:male ratio of D. tasmanica. Although coriander enhanced leafroller parasitism three-fold in field experiments compared with controls, it failed to enhance the longevity of both sexes of D. tasmanica in the laboratory compared with water-only. Broad bean significantly enhanced parasitoid abundance three-fold and significantly increased parasitism from 0% to 75% compared with the controls on one leafroller release date. However, laboratory trials showed that of male D. tasmancia but it did not enhance female longevity. Also, female D. tasmanica foraging on broad bean produced a total of only three parasitoid cocoons, but this result was based on an overall 6.5% survival of larvae to pupae or to parasitoid cocoon. Furthermore, results suggested that extrafloral nectar secretion decreased as the plants matured. Phacelia (Phacelia tanacetifolia Benth.) did not significantly enhance parasitism rate in the field compared with controls, and numbers of D. tasmanica captured by suction sampling were significantly lower in phacelia treatments compared with alyssum, buckwheat and control plots. Also, laboratory experiments showed that survival of D. tasmanica on phacelia flowers was equivalent to that on water-only and significantly lower than on buckwheat. These results suggest that phacelia does not provide nectar to D. tasmanica, only pollen, and therefore is not a suitable understorey plant for D. tasmanica enhancement in orchards. Buckwheat and alyssum showed the most potential as understorey plants for the enhancement of natural enemies. Buckwheat not only increased numbers of D. tasmanica seven-fold, but also increased numbers of beneficial lacewings (Micromus tasmaniae (Walker)) and hover flies (Syrphidae) captured on yellow sticky traps compared with the controls. It significantly increased leafroller parasitism by D. tasmanica from 0% to 86% compared with the controls (on one date only), and in the laboratory enhanced D. tasmanica longevity and increased fecundity compared with water-only. Similarly, alyssum significantly increased parasitism rate compared with controls, and two-fold more D. tasmanica were suction sampled in these plots compared with controls. It also enhanced longevity of both sexes of D. tasmanica compared with water, and showed the most favourable characteristics in terms of being of no benefit to leafrollers. This is because it was not preferred over apple by leafroller larvae and when they were forced to feed on it, it caused high mortality (94.3%) and low pupal weight (15 mg). Furthermore, alyssum did not enhance the number of fertile eggs produced by adult leafrollers compared with water only. However, further research is required to address the overall effect of buckwheat and alyssum on crop production and orchard management, including effects on fruit yield and quality, frost risk, disease incidence, soil quality, weeds and other pests. Also, research into the ability of these plants to survive in the orchard with little maintenance, and into the optimal sowing rates, would be useful. Sampling natural populations of leafroller within each treatment showed that damage from leafrollers and the number of leafroller larvae were respectively 20.3% and 29.3% lower in the flowering treatments compared with the controls. Furthermore, field trials showed up to a six-fold increase in leafroller pupae in controls compared with buckwheat and alyssum. This suggests that increasing leafroller parasitism rate from understorey management in orchards will translate into lower pest populations, although neither larval numbers/damage nor pupal numbers differed significantly between treatments. Trapping D. tasmanica at a gradient of distances showed that this parasitoid travels into rows adjacent to buckwheat plots, indicating that growers may be able to sow flowering plants in every second or third row of the orchard, and still enhance leafroller biocontrol while minimising the adverse effects of a cover crop. Sowing buckwheat and alyssum in orchard understoreys may enhance biological control of apple pests in organic apple production and reduce the number of insect growth regulators applied in IFP programmes. However, the challenge still remains to investigate whether conservation biological control can reduce leafroller populations below economic thresholds.

This study investigated understorey management in Canterbury, New Zealand, apple orchards for the enhancement of populations of Dolichogenidea tasmanica (Cameron) (Braconidae) for leafroller (Lepidoptera: Tortricidae) biological control. The first objective was to determine the influence of understorey plants on the abundance of D. tasmanica and leafroller parasitism, and to investigate the mechanisms behind this influence. The second was to determine the most suitable understorey plants in terms of their ability to enhance parasitoid abundance, leafroller parasitism, parasitoid longevity, parasitoid fecundity and its ability to not benefit leafroller. Results from three consecutive field trials showed that buckwheat (Fagopyrum esculentum Moench), coriander (Coriandrum sativum L.), alyssum (Lobularia maritima (L.) Desv), and, to a lesser extent, broad bean (Vicia faba L.), enhanced parasitoid abundance and leafroller parasitism. The mechanisms behind the effects of understorey plants had previously been unexplored. However, results here showed that it was the flowers or the buckwheat that 'attracted' the parasitoid to the plant and not the shelter, aphids or microclimate that the plant may also provide. Providing flowering plants in the orchard understorey also increased immigration of parasitoids and enhanced parasitoids and enhanced parasitoid longevity and fecundity in the laboratory. In contrast, the understorey plants had no influence on the female:male ratio of D. tasmanica. Although coriander enhanced leafroller parasitism three-fold in field experiments compared with controls, it failed to enhance the longevity of both sexes of D. tasmanica in the laboratory compared with water-only. Broad bean significantly enhanced parasitoid abundance three-fold and significantly increased parasitism from 0% to 75% compared with the controls on one leafroller release date. However, laboratory trials showed that of male D. tasmancia but it did not enhance female longevity. Also, female D. tasmanica foraging on broad bean produced a total of only three parasitoid cocoons, but this result was based on an overall 6.5% survival of larvae to pupae or to parasitoid cocoon. Furthermore, results suggested that extrafloral nectar secretion decreased as the plants matured. Phacelia (Phacelia tanacetifolia Benth.) did not significantly enhance parasitism rate in the field compared with controls, and numbers of D. tasmanica captured by suction sampling were significantly lower in phacelia treatments compared with alyssum, buckwheat and control plots. Also, laboratory experiments showed that survival of D. tasmanica on phacelia flowers was equivalent to that on water-only and significantly lower than on buckwheat. These results suggest that phacelia does not provide nectar to D. tasmanica, only pollen, and therefore is not a suitable understorey plant for D. tasmanica enhancement in orchards. Buckwheat and alyssum showed the most potential as understorey plants for the enhancement of natural enemies. Buckwheat not only increased numbers of D. tasmanica seven-fold, but also increased numbers of beneficial lacewings (Micromus tasmaniae (Walker)) and hover flies (Syrphidae) captured on yellow sticky traps compared with the controls. It significantly increased leafroller parasitism by D. tasmanica from 0% to 86% compared with the controls (on one date only), and in the laboratory enhanced D. tasmanica longevity and increased fecundity compared with water-only. Similarly, alyssum significantly increased parasitism rate compared with controls, and two-fold more D. tasmanica were suction sampled in these plots compared with controls. It also enhanced longevity of both sexes of D. tasmanica compared with water, and showed the most favourable characteristics in terms of being of no benefit to leafrollers. This is because it was not preferred over apple by leafroller larvae and when they were forced to feed on it, it caused high mortality (94.3%) and low pupal weight (15 mg). Furthermore, alyssum did not enhance the number of fertile eggs produced by adult leafrollers compared with water only. However, further research is required to address the overall effect of buckwheat and alyssum on crop production and orchard management, including effects on fruit yield and quality, frost risk, disease incidence, soil quality, weeds and other pests. Also, research into the ability of these plants to survive in the orchard with little maintenance, and into the optimal sowing rates, would be useful. Sampling natural populations of leafroller within each treatment showed that damage from leafrollers and the number of leafroller larvae were respectively 20.3% and 29.3% lower in the flowering treatments compared with the controls. Furthermore, field trials showed up to a six-fold increase in leafroller pupae in controls compared with buckwheat and alyssum. This suggests that increasing leafroller parasitism rate from understorey management in orchards will translate into lower pest populations, although neither larval numbers/damage nor pupal numbers differed significantly between treatments. Trapping D. tasmanica at a gradient of distances showed that this parasitoid travels into rows adjacent to buckwheat plots, indicating that growers may be able to sow flowering plants in every second or third row of the orchard, and still enhance leafroller biocontrol while minimising the adverse effects of a cover crop. Sowing buckwheat and alyssum in orchard understoreys may enhance biological control of apple pests in organic apple production and reduce the number of insect growth regulators applied in IFP programmes. However, the challenge still remains to investigate whether conservation biological control can reduce leafroller populations below economic thresholds.

Submitted by Jorge Silva (jorgelmsilva@ufrrj.br) on 2017-06-20T20:36:51Z No. of bitstreams: 1 2012 - Vinicius de Abreu D'?vila.pdf: 623654 bytes, checksum: 2c91585552193e53bcefc6b559fe2a2f (MD5) / Made available in DSpace on 2017-06-20T20:36:51Z (GMT). No. of bitstreams: 1 2012 - Vinicius de Abreu D'?vila.pdf: 623654 bytes, checksum: 2c91585552193e53bcefc6b559fe2a2f (MD5) Previous issue date: 2012-08-31 / CAPES / The biological control is as important method to regulate the pest populations in a system of sustainable agricultural production, because it is a promising alternative to the use of the organic synthetic pesticides that cause great ecotoxicological impacts. The predator ladybeetles are part of the biological control agents of agricultural pests, could be management by the three biocontrol strategies: classical, conservative and augmentative. In the present work, it was tried to generate knowledge for using the aphidophagous predator ladybeetle Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae) under the perspective of the last two strategies. The conservative biological control involving predator insects bases on the fact that in the absence or scarceness of their preferential prey or in the presence of the other preys with inferior nutritional quality, they may use alternative foods, such as pollen, to guarantee their survivorship and, sometimes, their reproduction, and because of that botanical species that provide this floral resource might integrate the agricultural landscape, inside and/or around the agricultural property; meanwhile the augmentative control requests the multiplication of the predator in the laboratory, using natural or artificial preys. Even though some authors proved the visitation of the flowers of some species of Apiaceae by C. maculata, there are no records in the literature of the ingestion of pollen grains of this botanical family by this ladybeetle. In this context, this work was carried out with the aim to select the plant species whose flowers are source of pollen as alternative or complementary food to C. maculata in the perspective to compose the vegetation of the agroecosystems to contribute in the conservation of this ladybeetle, and /or to aid in its mass rearing in the laboratory conditions. The objective of the chapter I was to prove the ingestion of pollen of three species of the family Apiaceae [coriander (Coriandrum sativum L.), dill (Anethum graveolens L.), and fennel (Foeniculum vulgare Mill.)] from the provision of their flowers to the larvae of the 4th instar and adults of C. maculata. It was observed the presence of pollen grains in the five replicates of all treatments, proving the ingestion of the pollen of these three species of Apiaceae from their flowers by C. maculata. At 24 hours of exposition, adults fed on average more pollen of dill than pollens of coriander and fennel, while the larvae consumed more pollen of fennel. The objective of the chapter II was to determine the suitability of nine diets to C. maculata, including provision of pollen of the two species of Apiaceae (coriander and dill), under controlled conditions of the laboratory. Even though the diets with only flowers of these two Apiaceae did not provided the full development of C. maculata, they used as complementary food with eggs of Anagasta kuehniella Zeller (Lepidoptera: Pyralidae) resulted in reduction of larval period, increased the egg number by cluster, and increased the body weight. The diet with alive larvae of Drosophila melanogaster Meigen (Diptera: Drosophilidae) was proved to be an essential food as well as resulted in adults with higher body weight, and the number of eggs per cluster increased in comparison with the feeding with only eggs of A. kuehniella. / O controle biol?gico ? um importante m?todo para regular as populac?es de pragas em um sistema de produ??o agr?cola sustent?vel, pois ? uma alternativa promissora ao uso de agrot?xicos org?nicos sint?ticos que causam grandes impactos ecotoxicol?gicos. As joaninhas predadoras fazem parte dos agentes de controle biol?gico de pragas agr?colas, podendo ser manejadas pelas tr?s estrat?gias de controle biol?gico: cl?ssico, conservativo e aumentativo. No presente trabalho, buscou-se gerar conhecimento para uso da joaninha predadora afid?faga Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae) sob a perspectiva das duas ?ltimas estrat?gias. O controle biol?gico conservativo envolvendo insetos predadores baseia-se no fato de que, na aus?ncia ou escassez da presa preferencial ou na presen?a de outras presas de qualidade inferior, podem usar alimentos alternativos, tais como p?len, para garantir sua sobreviv?ncia e, por vezes, sua reprodu??o e, por isso, esp?cies bot?nicas provedoras desse recurso floral devem integrar a paisagem agr?cola, dentro e/ou no entorno da propriedade agr?cola; enquanto o controle aumentativo requer a multiplica??o do predador no laborat?rio, podendo se valer de presas naturais ou artificiais. Apesar de alguns autores comprovarem a visita??o das flores de algumas esp?cies de Apiaceae por C. maculata, n?o h? relatos na literatura da ingest?o de gr?os de p?len dessa fam?lia bot?nica por essa joaninha. Nesse contexto, este trabalho foi conduzido a fim de selecionar esp?cies de plantas cujas flores sejam fonte de p?len como alimento alternativo ou complementar para C. maculata na perspectiva de compor a vegeta??o dos agroecossistemas para contribuir na conserva??o dessa joaninha, e/ou auxiliar na cria??o massal da mesma em condi??es de laborat?rio. O objetivo do cap?tulo I foi comprovar a ingest?o de p?len de tr?s esp?cies da fam?lia Apiaceae [coentro (Coriandrum sativum L.), endro (Anethum graveolens L.) e erva-doce (Foeniculum vulgare Mill.)] a partir da oferta de suas flores para larvas de 4? instar e adultos de C. maculata. Constatou-se a presen?a de gr?os de p?len nas cinco repeti??es de todos os tratamentos, comprovando a ingest?o de p?len dessas tr?s Apiaceae a partir de suas flores por C. maculata. Em 24 horas de exposi??o, os adultos consumiram em m?dia mais p?len de endro em compara??o aos polens de coentro e erva-doce, enquanto que as larvas consumiram mais p?len de erva-doce. O objetivo do capitulo II foi determinar a adequabilidade de nove dietas para C. maculata, incluindo oferta de p?len de duas esp?cies de Apiaceae (coentro e endro), em condi??es controladas de laborat?rio. Apesar das dietas apenas com flores dessas duas Apiaceae n?o proporcionarem o desenvolvimento completo de C. maculata, elas usadas com complementa??o da alimenta??o com ovos de Anagasta kuehniella Zeller (Lepidoptera: Pyralidae) possibilitam a redu??o do per?odo larval, aumento no n?mero de ovos por postura e aumento do peso corp?reo. A dieta com larvas vivas de Drosophila melanogaster Meigen (Diptera: Drosophilidae) n?o foi s? comprovada como alimento essencial como tamb?m resultou em adultos de maior peso corp?reo e um aumento no n?mero de ovos por postura em compara??o ? alimenta??o apenas com ovos de A. kuehniella.

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.