Thesis (MSc)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Invasive arthropod predators are one of the largest and most diverse groups of invasive insects in the world. Many are generalist predators, with cosmopolitan distributions due to their use as biological control agents in agriculture. Harmonia axyridis (Coleoptera: Coccinellidae), an invasive arthropod predator species native to Asia, which now has a world-wide distribution. It is considered one of the most successful biological control predator species and is generally considered to be economically beneficial. However, negative effects have recently emerged in agricultural and natural systems. Harmonia axyridis poses a threat to biodiversity as it outcompetes native species for food resources. It can also feed directly on native predatory arthropods that disrupt natural ecosystem processes. Their movement in-and-out of agricultural landscapes may depend on food availability with natural vegetation alongside agricultural areas often utilised for refuge and alternative food resources. This beetle has also been recorded in urban areas. The aim of this study was to determine how the invasive H. axyridis beetle uses the local landscape in the Western Cape province, South Africa, and to determine its threat to native species. I sampled urban landscapes, vineyards, natural vegetation/vineyard edge zones and pristine natural areas for arthropods every second month using a suction sampler. Data collected included the abundance and diversity of H. axyridis, herbivores, local predators and non-Harmonia ladybeetles. Most H. axyridis were collected in urban areas during all sampling periods. Highest abundance was recorded in May and July (winter). This indicates that urban areas were the preferred landscape feature and that these act as ovipositing areas, particularly as larval H. axyridis were also only collected in urban areas. Significantly, vineyards and natural vegetation had very low abundance of H. axyridis, questioning their value as a biological control agent in this region. Harmonia axyridis had a negative effect on the overall local arthropod community, as well as the predator and herbivore guilds, although it was positively correlated with the abundance of non-Harmonia ladybeetles. This suggests that H. axyridis and non-Harmonia ladybeetles are responding to the same resources in these landscapes. A negative correlation found between H. axyridis and the abundance of predators is most likely due to competition for the same resources (e.g. prey items). These negative impacts, along with their negligible value as biological control agents in agriculture, suggest that a programme should be implemented to control this invasive species. More specifically, control should be aimed in urban areas during winter when and where the species aggregates and when larvae are present. / AFRIKAANSE OPSOMMING: Geleedpotige roofdiere is een van die grootste en mees diverse groepe van uitheemse insekte in die wêreld. Die meeste is veelsydige roofdiere, met wêreldwye verspreiding te danke aan hul gebruik as biologiese beheer agente in landbou gebiede. Byvoorbeeld, Harmonia axyridis (Coleoptera: Coccinellidae), 'n indringer geleedpotige roofdier spesies inheems aan Asië, het nou 'n wêreldwye verspreiding. Dit word beskou as die mees suksesvolle roofdier spesies wat gebruik word vir biologiese beheer en word oor die algemeen beskou as ekonomies voordelig. Negatiewe effekte was onlangs aangeteken beide in landbou gebiede en natuurlike areas. Harmonia axyridis hou 'n bedreiging in vir inheemse biodiversiteit as dit inheemse spesies uitkompeteer vir voedsel bronne. Dit kan ook direk voed op plaaslike roofsugtige geleedpotiges wat trofiese vlakke ontwrig en uiteindelik, biodiversiteit. Hulle beweging in-en-uit landbou landskappe kan gekoppel word aan die beskikbaarheid van voedsel, en gebruik natuurlike plantegroei langs landbou gebiede dikwels as 'n toevlugs oord en area vir alternatiewe voedsel bronne. Harmonia axyridis word ook in stedelike gebiede aangeteken. Die doel van hierdie studie was om te bepaal hoe die indringer Harlekynkewer die plaaslike landskap gebruik met die fokus op wingerde in die Wes-Kaap provinsie van Suid-Afrika, en tweedens om die bedreiging wat hierdie kewer moontlik vir inheemse spesies te bepaal. Ek het arthropoda in stedelike landskappe, wingerde, natuurlike plantegroei / wingerd rand sones en ongerepte natuurlike areas elke twee maande met behulp van 'n D-vac versamel. Monsters was ontleed deur gebruik te maak van die getalle van H. axyridis, herbivore, plaaslike roofdiere en nie-Harmonia liewenheers kewers. Die meeste H. axyridis was in stedelike gebiede versamel gedurende al die seisoene, maar meeste individue was gedurende Mei en Julie (winter) versamel. Hierdie toon dat stedelike gebiede die voorkeur-landskappe is vir hierdie kewers en dat hierdie gebiede opgetree as eierleggende gebiede, veral omdat larwes van H. axyridis slegs in hierdie gebiede aangeteken was. Wingerde en die natuurlike plantegroei het baie lae getalle H. axyridis gehuisves wat hul waarde as biobeheermiddel bevraagteken. Harmonia axyridis het 'n negatiewe uitwerking op die algehele plaaslike geleedpotige gemeenskappe gehad, asook op die die roofdier en herbivoor gildes, maar hul getalle was positief gekorreleer met die getalle van nie-Harmonia liewenheerskewers. Dit dui daarop dat H. axyridis en nie-Harmonia liewenheerskewers beide reageer op dieselfde hulpbronne in hierdie landskappe. 'n Negatiewe korrelasie was gevind tussen die getalle van H. axyridis en die getalle van ander predatoriese geleedpotiges at waarskynlik te danke was aan mededinging tussen hierdie groepe vir dieselfde hulpbronne (bv prooi). Hierdie negatiewe invloede, asook hul verminderde waarde as biobeheeragente in die landbou, dui daarop dat 'n program in werking gestel moet word om hierdie indringerspesies te beheer. Meer spesifiek, beheer moet gedurende die winter en in stedelike gebiede geskied, waar en wanneer hierdie spesie op sy volopste is en waar larwes teenwoordig is.

Most Aphidinae species produce and use (E)-ß-farnesene (Eßf) as an alarm pheromone. This sesquiterpene is released by individuals under attack by a predator, and nearby aphids exhibit a variety of alarm behaviours. This PhD thesis aims to better understand how aphids manage their production and emission of alarm pheromone (Chapter IV). We also wanted, in a second step, to improve our knowledge on the roles that Eßf could play in the relationships that aphids have with their predators (Chapter V) and tending ants (Chapter VI), in order to better pinpoint the problem in this very tough context. The aphid predators have indeed a real advantage to be able to use the odorant cues emitted by their prey, to locate them and to select an adequate oviposition site. Ants establish with certain aphid species mutualistic relationships, which occurrence could be facilitated by the use of aphids odours. In Chapter IV, we have highlighted that aphid colonies non subjected to attack by predators release constantly small quantities of Eßf in their headspace, which means that this molecule could have additional roles than just acting as an alarm pheromone. In a second study, we demonstrated that the release of Eßf was not contagious, and therefore that a non stressed aphid receiving the alarm signal does not release additional Eßf. Since the production of alarm pheromone is likely to entail physiological cost, we tested and validated the hypothesis that aphids regulate their Eßf production according to their social environment. In Chapter V, we studied the ability of the hoverfly predator Episyrphus balteatus (Diptera, Syrphidae) to be used as biological control agent against aphids infesting tomato plants (Lycopersicon esculentum). After identifying the odours emitted by aphid infested plants, we have demonstrated that although this Diptera is able to perceive all the odours released by the system, it mainly uses Eßf to select its oviposition site. However, the E. balteatus larvae are not adapted to the architecture of tomato plants. We also showed that the Asian ladybeetle Harmonia axyridis (Coleoptera, Coccinellidae) olfaction was adapted to the perception of Eßf and that this beetle is also attracted by this sesquiterpene. Finally, in Chapter VI, we characterized the benefits accruing to aphid populations that have established mutualistic relationships with Lasius niger (Hymenoptera, Formicidae), and have demonstrated the role of Eßf and honeydew, respectively in locating aphid colonies and in the persistence of the mutualism. La plupart des espèces de pucerons appartenant à la sous-famille des Aphidinae produisent et utilisent le (E)-ß-farnésène (Eßf) comme phéromone dalarme. Ce sesquiterpène est relargué par les individus stressés par lattaque dun prédateur et cause chez les individus qui le perçoivent un comportement dalerte. La présente thèse de doctorat a pour objectif de comprendre comment les pucerons gèrent la production et émission de phéromones dalarme (Chapitre IV). Nous voulions aussi, dans une seconde étape, améliorer les connaissances sur les rôles potentiels que cet Eßf peut jouer au sein des relations que les pucerons entretiennent avec leurs prédateurs (Chapitre V) et avec les fourmis (Chapitre VI) Les prédateurs de pucerons retirent en effet un réel avantage à pouvoir saider des odeurs émises par leurs proies pour les localiser et pour sélectionner un site doviposition adéquat. Les fourmis, quant à elles, établissent des relations de mutualisme avec certaines espèces de pucerons. Les rencontres entre fourmis et pucerons pourraient être facilitées par lutilisation des odeurs de pucerons. Les résultats obtenus peuvent être résumés de la manière suivante: Dans le chapitre IV, nous avons mis en évidence que les colonies de pucerons non soumises à lattaque de prédateurs relarguent constamment de faibles quantités dEßf, ce qui permet dassumer que cette molécule puisse avoir dautres fonctions que celle de phéromone dalarme. Dans une deuxième étude, nous avons démontré que lémission du Eßf nétait pas contagieuse, et donc quun puceron non stressé percevant le signal dalarme német pas à son tour de le Eßf. Puisque la production de phéromone dalarme a inévitablement un coût physiologique, nous avons testé et validé lhypothèse selon laquelle les pucerons régulent leur production de Eßf en fonction de leur environnement social. Dans le chapitre V, nous avons étudié la possibilité dutiliser le syrphe prédateur Episyrphus balteatus (Diptera, Syrphidae) en lutte biologique contre les pucerons infestant les plants de tomate (Lycopersicon esculentum). Après avoir identifié les odeurs émises par les plants infestés, nous avons démontré que, si ce Diptère est capable de percevoir lensemble des odeurs émises par ce système tritrophique, il utilise principalement lEßf pour sélectionner son site doviposition. Cependant, les larves dE. balteatus ne sont pas adaptées à larchitecture des plants de tomate. Nous avons également montré que la coccinelle asiatique Harmonia axyridis (Coleoptera, Coccinellidae) possède le matériel olfactif nécessaire à la perception du Eßf et quelle est aussi attirée par ce sesquiterpène. Enfin, dans le chapitre VI, nous avons caractérisé les bénéfices retirés par les populations de pucerons ayant établi des relations de mutualisme avec Lasius niger (Hymenoptera, Formicidae), et avons démontré le rôle du Eßf et du miellat, respectivement dans la localisation des colonies de pucerons et dans la persistance du mutualisme.

Fitness of insecticide resistant insect populations can be different to susceptible insect populations regarding adaptive costs. Therefore, this study investigated the fitness of Eriopis connexa (Germar) (Coleoptera: Coccnellidae) population resistant to lambda-cyhalothrin, as well as its predation rate on Aphis gossypi (Glover) (Hemiptera: Aphididae). The insects used to run the bioassays were reared under laboratory conditions of 25 ± 1oC and 12:12h (L:D) photoperiod and fed with Anagasta kuehniella (Zeller) (Lepidoptera: Pyralidae) eggs. The field resistant population of E. connexa (RR) was maintained under selection pressure at each adult generation in the laboratory by exposing them to the LD50-dose (0.25 mg a.i./mL of lambda-cyhalothrin at technical grade) determined after 12 successive generations of selections in the laboratory. The susceptible population (SS) was reared at the same condition; however, without exposing the insects to the insecticide. Non-treated RR females and treated recovered females (RRr) after lambda-cyhalothrin topic treatment with 0.05, 0.10 and 0.25 mg a.i./mL produced on average 50% less eggs than SS females. All other adult life history characteristics were similar among SS, RR and RRr populations. Among the developmental characteristics for offspring produced by SS, RR, and RRr females was observed lower larval viability and weight of adult males for RRr offspring. Furthermore, females from SS, RR and RRr populations exposed to prey shortage between 3rd- and 13th-day in the adult stage exhibited statistically similar survival ranging from 48 to 63 days. However, egg production was significantly higher for females SS (average = 290 eggs), followed by females RR (180 eggs), and RRr (50 eggs) during this period. The average consumption of cotton aphid during five consecutive days was significantly higher for ladybeetles SS, followed by RR and RRr up to the third day of observation. However, after the fourth day, there was no difference in the aphid consumption among the three populations SS, RR, and RRr. The results show that females RR have lower reproductive output in comparison to females SS, and it is not related to the knockdown effect; however, the costs of recovering from knockdown interferes with egg production under food shortage, with larval viability and adults predation rate. Thus, we can conclude that E. connexa population resistant to the lambda-cyhalothrin exhibits adaptive costs which is increased when subjected to prey shortage. / Conselho Nacional de Desenvolvimento Científico e Tecnológico / O desempenho de populações de insetos resistentes a inseticidas pode diferir daquelas susceptíveis caracterizando um custo adaptativo. Assim, este estudo investigou o desempenho de uma população resistente de Eriopis connexa (Germar) (Coleoptera: Coccinellidae) à lambda-cialotrina, bem como sua taxa de predação sobre Aphis gossypii (Glover) (Hemiptera: Aphididae). Os insetos resistentes empregados nos estudos foram criados em laboratório a 25 ± 1oC, fotoperíodo de 12:12h (L:E) e alimentados com ovos de Anagasta kuehniella (Zeller) (Lepidoptera: Pyralidae). A população de campo de E. connexa resistente (RR) foi mantida sob pressão de seleção com tratamento tópico do adulto com 0,5 μl da concentração 0,25 mg i.a./mL de lambda-cialotrina em grau técnico correspondente a DL50 determinada na 12a geração após sucessivas seleções em laboratório. A população susceptível (SS) foi criada sob as mesmas condições sem exposição ao inseticida. Fêmeas RR não tratadas e fêmeas RR recuperadas (RRr) do knockdown após tratamento tópico com lambda-cialotrina nas concentrações 0,05; 0,10 e 0,25 mg de i.a./mL produziram em média 50% menos ovos, comparada as fêmeas SS independente da concentração usada do inseticida. As demais características da fase adulta não diferiram entre fêmeas SS, RR e RRr. Entre as características do desenvolvimento da progênie oriunda de fêmeas SS, RR e RRr observou-se menor viabilidade de larvas e peso de machos para descendentes de fêmeas RRr. Fêmeas submetidas à condição de escassez de presas do 3o ao 13o dia de vida apresentaram sobrevivência similar entre as populações SS, RR e RRr variando de 48 a 63 dias. Porém, a produção de ovos foi maior para fêmeas SS (290 ovos), seguida de fêmeas RR (180 ovos) e RRr (50 ovos) neste período. O consumo médio de pulgões durante cinco dias de observação foi significativamente superior para joaninhas SS que joaninhas RR e RRr, comportando-se desta maneira até o terceiro dia de observação. A partir do quarto dia, no entanto, não é detectada diferença na taxa diária de predação de pulgão entre as três populações SS, RR e RRr. Os resultados mostraram que fêmeas RR apresentam menor desempenho reprodutivo comparado a fêmeas SS, independente da ação knockdown, mas que o custo da recuperação do knockdown interfere na produção de ovos após escassez de alimento, na viabilidade larval e na taxa de predação. Estes resultados mostram que E. connexa possui um custo devido à resistência à lambda-cialotrina, e que o mesmo é aumentado em situações de escassez de presas.

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.