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During the past decade, the use of predatory mirids alone or combined with releases of egg parasitoids of the genus Trichogramma have been tested in Europe for biological control of the worldwide invasive pest, Tuta absoluta (Meyrick). Here, we evaluated the control of this pest by the release of the Neotropical mirid Macrolophus basicornis (Stal), the Neotropic/Nearctic parasitoid Trichogramma pretiosum Riley, and by combined releases of the predator and the parasitoid. Tests were conducted in greenhouse compartments during the summer and fall season. Each compartment contained 10 tomato plants in which only the pest was released or the pest with 1 or 2 natural enemies. Plant damage, and pest and natural enemy densities were checked weekly on one apical, medium, and bottom leaf of 5 plants. Both M. basicornis and T. pretiosum significantly reduced T. absoluta density when released alone. Combined releases resulted in a 10% higher reduction during the summer season, but not during the fall season. The damage caused by T. absoluta was significantly higher in control treatments than in all natural enemy treatments: at the end of the summer trial leaves were completely damaged in the control treatment, whereas only up to 25% leaf damage occurred in the natural enemy treatments. Combined releases did not result in lower damage than with releases of either M. basicornis or T. pretiosum. Practical aspects of single and combined releases are discussed.

Biological control, which uses natural enemies to reduce pest populations, is a non-polluting powerful method to manage impacts of (invasive) pests. Currently, polyphagous mirid predators are increasingly used in commercial, augmentative biological control. Information about their foraging behaviour is essential, especially if one intends to use several natural enemies for control of one or more pests in a crop, to detect if negative intraguild effects occur. We studied a case of intraguild predation (IGP) involving a predator, Macrolophus basicornis, of the worldwide invasive South American tomato leaf miner Tuta absoluta, and explored how this predator deals with prey parasitized by Trichogramma pretiosum. Behavioural observations show that M. basicornis predators contacted significantly fewer old, parasitized eggs of T. absoluta than recently parasitized ones. Olfactometer tests revealed that predators could smell differences between volatiles of tomato leaves infested with eggs of different quality to locate suitable prey. They preferred volatiles from leaflets with unparasitized eggs above control leaflets, and, moreover, preferred volatiles from leaflets with recently parasitized eggs over volatiles of leaflets with 5-day old parasitized eggs. When predators and parasitoids are used together to control T. absoluta, parasitoids should be introduced days before predators to prevent high levels of IGP.

Environmental risk assessments (ERAs) are required before utilizing exotic arthropods for biological control (BC). Present ERAs focus on exposure analysis (host/prey range) and have resulted in approval of many specialist exotic biological control agents (BCA). In comparison to specialists, generalist arthropod BCAs (GABCAs) have been considered inherently risky and less used in classical biological control. To safely consider exotic GABCAs, an ERA must include methods for the analysis of potential effects. A panel of 47 experts from 14 countries discussed, in six online forums over 12 months, scientific criteria for an ERA for exotic GABCAs. Using four case studies, a three-tiered ERA comprising Scoping, Screening and Definitive Assessments was developed. The ERA is primarily based on expert consultation, with decision processes in each tier that lead to the approval of the petition or the subsequent tier. In the Scoping Assessment, likelihood of establishment (for augmentative BC), and potential effect(s) are qualitatively assessed. If risks are identified, the Screening Assessment is conducted, in which 19 categories of effects (adverse and beneficial) are quantified. If a risk exceeds the proposed risk threshold in any of these categories, the analysis moves to the Definitive Assessment to identify potential non-target species in the respective category(ies). When at least one potential non-target species is at significant risk, long-term and indirect ecosystem risks must be quantified with actual data or the petition for release can be dismissed or withdrawn. The proposed ERA should contribute to the development of safe pathways for the use of low risk GABCAs.

Carabid beetles are common predators of pest insects and weed seeds in agricultural systems. Understanding their dispersal across farmland is important for designing farms and landscapes that support pest and weed biological control. Little is known, however, about the effect of farmland habitat discontinuities on dispersal behaviour and the resulting redistribution of these beetles. We released 1,985 well-fed and 1,680 food-deprived individuals of the predatory carabid beetle Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on a farm in Wageningen, The Netherlands. We recaptured 23.6% of those beetles over a period of 23 days in 2010. The farmland comprised agricultural fields with various crop species and tillage, separated by strips of perennial vegetation. We developed discrete Fokker-Planck diffusion models to describe dispersal based on motility (m2 day−1) and preferential behaviour at habitat interfaces. We used model selection and Akaike’s information criterion to determine whether movement patterns were driven by variation in motility between habitats, preferential behaviour at habitat interfaces, or both. Model selection revealed differences in motility among habitats and gave strong support for preferential behaviour at habitat interfaces. Behaviour at interfaces between crop and perennial vegetation was asymmetric, with beetles preferentially moving towards the crop. Furthermore, beetles had lower motility in perennial strips than in arable fields. Also between arable habitats movement was asymmetric, with beetles preferentially moving towards the habitat in which motility was lowest. Neither crop type nor tillage explained differences in motility between crop habitats. Recapture data representing dispersal patterns of beetles were best described by a model that accounted for differences in motility between farmland habitats and preferential behaviour at habitat interfaces. Motility in farmland and behaviour at interfaces can also be estimated for other organisms and farmland habitats to support design of farmland conducive to natural pest suppression. Landscape design for early recruitment of carabids into arable fields should take into account the quantity and quality of resource habitats in the landscape, their proximity to crop fields, movement rates, and the possibility of movement responses at interfaces between landscape elements.

Plants with glandular trichomes may provide protection against herbivores by impeding their movement, but may also hinder natural enemies. We investigated walking behavior and predation rates of the mirid predators Campyloneuropsis infumatus, Engytatus varians, and Macrolophus basicornis on two tomato lines with different densities of sticky trichomes. Also, the time spent by each predator species in various activities (immobile, walking, grooming, probing, and feeding) and the searching for and handling time of prey, eggs of Tuta absoluta, were observed. Tomato lines TOM 587 and TOM 687 were used, with TOM 687 having a 179% higher density of glandular trichomes. All three mirid predators easily climbed the sticky stems of both tomato lines and found eggs of T. absoluta on leaflets located on the top of the plants. Predation rates, grooming, probing, and feeding activities and the time spent by the three mirid predators until prey encounter were not influenced by a difference in density of trichomes on the two tomato lines. However, E. varians walked longer on TOM 584, and spent more time grooming than M. basicornis. Macrolophus basicornis walked longer on TOM 687. Our results show that the three mirid predators are apparently specialists of glandular plants, because differences in trichome density do not influence their activities, including their prey finding efficiency.

Due to typesetting errors, Table 5 was not displayed correctly in the initial online publication. The original online article has been corrected.

In response to herbivory by insects, various plants produce volatiles that attract enemies of the herbivores. Although ants are important components of natural and agro-ecosystems, the importance of herbivore-induced plant volatiles (HIPVs) as cues for ants for finding food sources have received little attention. We investigated responses of the ant Formica pratensis to volatiles emitted by uninfested and insect-infested cucumber (Cucumis sativus) and potato (Solanum tuberosum) plants. Cucumber plants were infested by the phloem-feeding aphid Aphis gossypii, the leaf chewer Mamestra brassicae or simultaneously by both insects. Potato plants were infested by either Aphis gossypii, by the leaf chewer Chrysodeixis chalcites or both. In olfactometer experiments, ants preferred volatile blends emitted by cucumber plants infested with M. brassicae caterpillars alone or combined with A. gossypii to volatiles of undamaged plants or plants damaged by A. gossypii only. No preference was recorded in choice tests between volatiles released by aphid-infested plants over undamaged plants. Volatiles emitted by potato plants infested by either C. chalcites or A. gossypii were preferred by ants over volatiles released by undamaged plants. Ants did not discriminate between potato plants infested with aphids and caterpillars over plants infested with aphids only. Plant headspace composition showed qualitative and/or quantitative differences between herbivore treatments. Multivariate analysis revealed clear separation between uninfested and infested plants and among herbivore treatments. The importance of HIPVs in indirect plant defence by ants is discussed in the context of the ecology of ant-plant interactions and possible roles of ants in pest management.

We consider IPM as a combination of durable, environmentally, toxicologically and economically justifiable farming practices which prevent pest damage primarily through the use of natural factors limiting pest population growth and disease development, and which resort only if needed to other, preferably non-chemical, measures. IPM is not simply a combination of various control methods. We give an overview of IPM measures used in greenhouses and refer to specific chapters in this book for examples. In IPM, each practical situation dictates a number of special aspects for consideration, and IPM methods need continuous adaptation, making IPM knowledge intensive and interactive. Successful IPM programmes for greenhouse crops have a number of characteristics in common: (a) their use was promoted only after a complete IPM programme had been developed, (b) intensive support by the extension service was essential during initial implementation, (c) the costs of crop protection with IPM should not be higher than those of a chemical control programme, and (d) non-chemical management methods, such as biocontrol agents and resistant plant material, should be as easily available, as reliable, and as constant in quality, as chemical agents. IPM research and implementation in greenhouses during the past 50 years has taught us the lesson that the development of an IPM programme needs to be discussed in a very early stage with all stakeholders, including growers, pest management specialists, extension services and researchers. Such a meeting often results in a pragmatic design of a draft, very pragmatic IPM programme, which is continuously adapted during later meetings, based on growers’ experience and new research results.

Biocontrol on Haiti is mainly in the form of classical biological control. Successful early projects are control of citrus blackfly by a hymenopteran parasitoid since the 1930s and control of the sugarcane borer by an introduced tachinid parasitoid since 1953. After 1970 three hymenopteran parasitoids were introduced for control of the coffee berry borer. A new project concerns the introduction of predators against the fluted scale Crypticerya genistae. Most Haitian farmers have insufficient financial resources to apply chemical control, which creates possibilities for use of conservation biocontrol and also allows the ecosystem service of biocontrol to exhibit its role in pest and disease reduction.

First we describe the different types of biocontrol used in greenhouses and present examples of each type. Next we summarize the history of greenhouse biocontrol, which started in 1926, showed a problematic period when synthetic chemical pesticides became available after 1945, and flourished again since the 1970s. After 1970, the number of natural enemies becoming available for commercial augmentative biocontrol in greenhouses grew very fast, as well as the industry producting these control agents. Biocontrol of the most important clusters of greenhouse pests is summarized, as well as the taxonomic groups of natural enemies that play a main role in greenhouses. More than 90% of natural enemy species used in greenhouses belong to the Arthropoda and less than 10%, many belonging to the Nematoda, are non-arthropods. This is followed by sections on finding and evaluation of potential biocontrol agents, and on mass production, storage, release and quality control of natural enemies. Since the 1970s, production of biocontrol agents has moved from a cottage industry to professional research and production facilities. Many efficient agents have been identified, quality control protocols, mass-production, shipment and release methods matured, and adequate guidance for farmers has been developed. Most natural enemy species (75%) are produced in low or medium numbers per week (hundreds to a hundred thousand), and are applied in situations where only low numbers are needed, such as private gardens, hospitals, banks, and shopping malls. The other 25% of the species are produced in numbers of 100,000 to up to millions per week and regularly released in many of the greenhouse crops. Microbial pesticides are predominantly used as corrective treatments in greenhouse crops where natural enemies are providing insufficient control. Europe is still the largest commercial market for arthropod greenhouse biocontrol agents, and North America is the largest market for microbial control agents. We then continue with a discussion on the pros and cons of use of polyphagous predators, and the use of semiochemicals. Finally, we summarize factors that indicate a positive future for greenhouse biocontrol, as well as developments frustrating its implementation.

Tuta absoluta Meyrick quickly developed into a significant pest of tomatoes worldwide. While the mirid bugs Macrolophus basicornis (Stal), Engytatus varians (Distant) and Campyloneuropsis infumatus (Carvalho) prey on this tomato borer, their biology have not been well characterized. Using a mixture of T. absoluta eggs and larvae as food on tomato, mirid life histories were studied. Nymphs and adults successfully used T. absoluta as prey, although significant differences in biology of the species were observed. Campyloneuropsis infumatus and M. basicornis were quickest and slowest to become adult at 16 and 19 days, respectively. Nymphal survival was high and varied from 70 to 75 %. Female biased sex ratios were observed for C. infumatus (0.80) and M. basicornis (0.71), but not for E. varians (0.56). Intrinsic rates of increase for C. infumatus, E. varians and M. basicornis were 0.11, 0.10 and 0.11 respectively. Our results indicate rapid development and population increase of these three mirids, making them interesting candidates for biological control of T. absoluta on tomato.

Augmentative biological control uses mass-reared natural enemies for releases in large numbers to reduce pest populations. Augmentative biocontrol is increasingly applied worldwide in many different crops. It is a cornerstone of integrated pest management (IPM) programmes and plays a key sustainability-determining role in IPM. This chapter describes the scale of augmentative biological control and where this type of biocontrol is applied. The chapter describes the natural enemies commercially available and gives information on their mass production. The chapter discusses the important role of biocontrol in IPM programmes, and its practical use in very different agricultural production systems. Finally, the chapter presents the advantages and disadvantages of this control method, and looks ahead to future research trends in this area.

In augmentative biological control (ABC), invertebrate and microbial organisms are seasonally released in large numbers to reduce pests. Today it is applied on more than 30 million ha worldwide. Europe is the largest commercial market for invertebrate biological control agents, while North America has the largest sales of microbials. A strong growth in use of ABC, particularly of microbial agents, is taking place in Latin America, followed by Asia. The current popularity of ABC is due to (1) its inherent positive characteristics (healthier for farm workers and persons living in farming communities, no harvesting interval or waiting period after release of agents, sustainable as there is no development of resistance against arthropod natural enemies, no phytotoxic damage to plants, better yields and a healthier product, reduced pesticide residues [well below the legal Maximum Residue Levels (MRLs)], (2) professionalism of the biological control industry (inexpensive large scale mass production, proper quality control, efficient packaging, distribution and release methods, and availability of many (>440 species) control agents for numerous pests), (3) a number of recent successes showing how biological control can save agricultural production when pesticides fail or are not available, (4) several non-governmental organizations (NGOs), consumers, and retailers demanding pesticide residues far below the legal MRLs, and (5) policy developments in several regions of the world aimed at reduction and replacement of synthetic pesticides by more sustainable methods of pest management. We are convinced, however, that ABC can be applied on a much larger area than it is today. We plead in the short term for a pragmatic form of agriculture that is adaptable, non-dogmatic and combines the sustainability gain from all types of agriculture and pest management methods. We then propose to move to “conscious agriculture”, which involves participation of all stakeholders in the production and consumer chain, and respects the environment and resource availability for future generations. Were “conscious agriculture” to be considered a serious alternative to conventional farming, ABC would face an even brighter future.

Tuta absoluta (Meyrick; Lepidoptera: Gelechiidae) is a key pest of tomato and is quickly spreading over the world. We conducted an experiment aimed at evaluating the control capacity and risk for plant damage of three Neotropical mirid species, Campyloneuropsis infumatus (Carvalho; Hemiptera: Miridae), Engytatus varians (Distant; Hemiptera: Miridae) and Macrolophus basicornis (Stal; Hemiptera: Miridae) on T. absoluta infested tomato plants in large cages in an experimental greenhouse. During three successive periods of 9 wk each, we followed population development of the three mirids when exposed to T. absoluta, and of T. absoluta alone in separate cages in the greenhouse. We determined weekly the numbers of T. absoluta eggs and larvae per leaf, the number of mirid predators per leaf, the percentage of damaged leaves and fruits by T. absoluta, and the weight of fruits. Two of the mirid predators, C. infumatus and M. basicornis, successfully established on T. absoluta infested tomato plants and significantly reduced T. absoluta numbers, which ultimately resulted in an increased yield. These two mirid species hardly injured tomato plants or fruits as a result of plant feeding. Surprisingly, the species E. varians, which showed high predation rates in laboratory experiments, did not establish and reduce pest populations in any of the tests.

Plants emit volatile compounds in response to insect herbivory, which may play multiple roles as defensive compounds and mediators of interactions with other plants, microorganisms and animals. Herbivore-induced plant volatiles (HIPVs) may act as indirect plant defenses by attracting natural enemies of the attacking herbivore. We report here the first evidence of the attraction of three Neotropical mirid predators (Macrolophus basicornis, Engytatus varians and Campyloneuropsis infumatus) toward plants emitting volatiles induced upon feeding by two tomato pests, the leaf miner Tuta absoluta and the phloem feeder Bemisia tabaci, in olfactometer bioassays. Subsequently, we compared the composition of volatile blends emitted by insect-infested tomato plants by collecting headspace samples and analyzing them with GC-FID and GC-MS. Egg deposition by T. absoluta did not make tomato plants more attractive to the mirid predators than uninfested tomato plants. Macrolophus basicornis is attracted to tomato plants infested with either T. absoluta larvae or by a mixture of B. tabaci eggs, nymphs and adults. Engytatus varians and C. infumatus responded to volatile blends released by tomato plants infested with T. absoluta larvae over uninfested plants. Also, multiple herbivory by T. absoluta and B. tabaci did not increase the attraction of the mirids compared to infestation with T. absoluta alone. Terpenoids represented the most important class of compounds in the volatile blends and there were significant differences between the volatile blends emitted by tomato plants in response to attack by T. absoluta, B. tabaci, or by both insects. We, therefore, conclude that all three mirids use tomato plant volatiles to find T. absoluta larvae. Multiple herbivory did neither increase, nor decrease attraction of C. infumatus, E. varians and M. basicornis. By breeding for higher rates of emission of selected terpenes, increased attractiveness of tomato plants to natural enemies may improve the effectiveness of biological control.

Landscape structure as well as local vegetation influence biodiversity in agroecosystems. A study was performed to evaluate the effect of floristic diversity, vegetation patterns, and landscape structural connectivity on butterflies (Lepidoptera: Papilionoidea and Hesperiidae), carabids (Coleoptera: Carabidae), syrphids (Diptera: Syrphidae), and sawflies (Hymenoptera: Symphyta). Vegetation analysis and insect samplings were carried out in nine sites within an intensively farmed landscape in northern Italy. Plant species richness and the percentage of tree, shrub, and herb cover were determined by means of the phytosociological method of Braun-Blanquet. Landscape structural connectivity was measured as the total length of hedgerow network (LHN) in a radius of 500 m around the center of each sampling transect. Butterflies species richness and abundance were positively associated both to herb cover and to plant species richness, but responded negatively to tree and shrub cover. Shrub cover was strictly correlated to both species richness and activity density of carabids. The species richness of syrphids was positively influenced by herb cover and plant richness, whereas their abundance was dependent on ligneous vegetation and LHN. Rarefaction analysis revealed that sawfly sampling was not robust and no relationship could be drawn with either vegetation parameters or structural connectivity. The specific responses of each insect group to the environmental factors should be considered in order to refine and optimize landscape management interventions targeting specific conservation endpoints.

Knowledge about the orientation mechanisms used by two important predaceous mirids (Macrolophus pygmaeus Rambour and Nesidiocoris tenuis (Reuter)) in finding their prey (whitefly Bemisia tabaci (Gennadius) and the tomato borer Tuta absoluta (Meyrick)) is limited. In a Y-tube olfactometer, we tested the behavioral responses of naïve and experienced predators to uninfested plants, herbivore-induced plant volatiles (HIPVs) from plants infested with T. absoluta and/or B. tabaci, the sex pheromone of T. absoluta, and volatiles produced by plants injured by the predators. Nesidiocoris tenuis responds to volatiles produced by uninfested plants only after experience with the plant, whereas naïve and experienced M. pygmaeus show positive chemotaxis. Both predators are attracted to volatiles from prey-infested plants, and we provide the first evidence that experience affects this response in M. pygmaeus. Infestation of the same plant by both prey species elicited similar responses by the two predators as plants infested by either herbivore singly. Neither predator responded to sex pheromones of T. absoluta. Macrolophus pygmaeus avoided plants injured by conspecifics, while N. tenuis females were attracted by such plants. The implications of these results for augmentative biological control are discussed.

The harlequin ladybird, Harmonia axyridis, is an important natural enemy of aphids throughout the world, but is now also considered an invasive alien species. We performed a meta-analysis of published life history data to address the question whether invading populations in Europe and North America have life history parameters that differ from native populations in Asia, explaining the beetle’s invasion success in new territories. In this meta-analysis, we accounted for important covariables that are often reported in published studies such as temperature, food source (aphids or eggs of Ephestia kuehniella), strain (laboratory or field populations) and photoperiod. Temperature was a key factor having consistent large effects on development rate, survival and reproductive characteristics of H. axyridis. Food source, strain, and photoperiod had effects on some, but not all characteristics, and their overall effect across characteristics was minor. Individuals of invasive populations had a shorter pre-oviposition period and higher fecundity at low temperatures than those of native populations, and a greater longevity across all temperatures. No differences in survival were found between native and invasive populations, while differences in development rate were not consistent, with opposing results obtained according to the way development rate was measured in trials reported in the literature. Results of this meta-analysis support the hypothesis that the life history of the beetle has changed during its invasion into North America and Europe. Invasive populations had a shorter pre-oviposition period and higher fecundity at low temperatures, as well as a greater longevity across all temperatures than native populations. These differences may partially explain the invasive success of H. axyridis.

Will the Convention on Biological Diversity put an end to biological control? Under the Convention on Biological Diversity countries have sovereign rights over their genetic resources. Agreements governing the access to these resources and the sharing of the benefits arising from their use need to be established between involved parties. This also applies to species collected for potential use in biological control. Recent applications of access and benefit sharing principles have already made it difficult or impossible to collect and export natural enemies for biological control research in several countries. If such an approach is widely applied it would impede this very successful and environmentally safe pest management method based on the use of biological diversity. The International Organization for Biological Control of Noxious Animals and Plants has, therefore, created the "Commission on Biological Control and Access and Benefit Sharing". This commission is carrying out national and international activities to make clear how a benefit sharing regime might seriously frustrate the future of biological control. In addition, the IOBC Commission members published information on current regulations and perceptions concerning exploration for natural enemies and drafted some 30 case studies selected to illustrate a variety of points relevant to access and benefit sharing. In this article, we summarize our concern about the effects of access and benefit sharing systems on the future of biological control. Poderá a Convenção em Diversidade Biológica por um fim no Controle Biológico? Baseando-se na Convenção sobre Diversidade Biológica, os países têm soberania sobre seus recursos genéticos. Acordos que governam o acesso a tais recursos e o compartilhamento dos benefícios provenientes do seu uso precisam ser estabelecidos de comum acordo com as partes envolvidas. Isto também é aplicável a espécies coletadas com uso potencial em controle biológico. Recentes aplicações dos princípios de introdução e compartilhamento dos benefícios têm tornado difícil, ou mesmo impossível, coletar e exportar inimigos naturais em muitos paises para pesquisas em controle biológico em muitos países. Como esta é uma medida amplamente utilizada, tais procedimentos poderão impedir este bem sucedido e ambientalmente seguro método de manejo de pragas, baseado no uso da diversidade biológica. A Organização Internacional para Controle Biológico de Plantas e Animais Nocivos (IOBC) criou a "Comissão em Controle Biológico e Introdução e Benefícios Mútuos" para estudar o assunto. Tal comissão está desenvolvendo atividades nacionais e internacionais para esclarecer como o regime de compartilhamento de benefícios pode prejudicar seriamente o futuro do controle biológico. Além disto, membros da Comissão da IOBC publicaram informações sobre regulamentos atuais e suas opiniões relacionadas à exploração de inimigos naturais, listando cerca de 30 casos para ilustrar os pontos relevantes para a introdução e compartilhamento de benefícios. No presente artigo, é sumarizado o ponto de vista dessa comissão na IOBC sobre os efeitos dos sistemas de introdução e compartilhamento para o futuro do Controle Biológico.

Under the Convention on Biological Diversity (CBD) countries have sovereign rights over their genetic resources. Agreements governing the access to these resources and the sharing of the benefits arising from their use need to be established between involved parties [i.e. Access and Benefit Sharing (ABS)]. This also applies to species collected for potential use in biological control. Recent applications of CBD principles have already made it difficult or impossible to collect and export natural enemies for biological control research in several countries. If such an approach is widely applied it would impede this very successful and environmentally safe pest management method based on the use of biological diversity. The CBD is required to agree a comprehensive Access and Benefit Sharing process in 2010, in preparation for which the IOBC (International Organization for Biological Control of Noxious Animals and Plants) Global Commission on Biological Control and Access and Benefit Sharing has prepared this position paper. Here, we first describe the practice of biological control in relation to the principles of ABS, illustrated extensively by case studies and successes obtained with biological control. Next, we emphasise the very limited monetary benefits generated in biological control when compared to other fields of ABS such as the collection of germplasm for development of human drugs, chemical pesticides or crop cultivars. Subsequently, we inform the biological control community of good ABS practice and challenges, and we hope to make clear to the community involved in ABS under the CBD the special situation with regard to biological control. Finally, based on the non-commercial academic research model, we make recommendations which would facilitate the practice of collection and exchange of biological control agents, propose a workable framework to assist policy makers and biological control practitioners, and urge biological control leaders in each country to get involved in the discussions with their national ABS contact point to take their needs into consideration.

During 16 weeks absolute counts and random samples were carried out in a glasshouse to determine population density and distribution pattern of the greenhouse whitefly (Trialeurodes vaporariorum) and its parasite Encarsia formosa. The results obtained by both methods were compared, and it was concluded that the data from the random sampling reflected the actual numbers and the distribution of the whitefly and its parasite very badly. The results are too bad to recommend this method to growers for obtaining a reliable idea about the sites and the degree of an infestation on which they can base the number and the dates of parasite introductions. Zusammenfassung Zur Parasit-Wirt-Beziehung zwischen Encarsia formosa (Hym. Aphelinidae) und Trialeurodes vaporariorum (Homopt. Aleyrodidae). VIII. Vergleich und Bewertung zweier verschiedener Ermittlungen der Populationsdichte und des Verteilungsbildes Wahrend 16 Wochen wurden die Populationsdichte und das Verteilungsbild der Weisen Fliege (Trialeurodes vaporariorum) und ihres Parasiten (Encarsia formosa) in einem Gewachshaus unter Verwendung von zwei Methoden ermittelt: eine unmittelbare Zahlung aller Individuen und eine zufallige Stichprobennahme. Die Ergebnisse beider Methoden wurden verglichen. Es zeigte sich, das die Daten der zufalligen Stichproben die wirkliche Abundanz und das Verteilungsbild der Weisen Fliege und ihres Parasiten nur sehr schlecht wiedergaben. Um zuverlassige Daten uber die Lokalisation und Grose der verschiedenen Herden zu bekommen, auf denen die Zuchter die Anzahl und die Zeitpunkte der Parasiteneinfuhrungen grunden konnen, sollte man daher die Stichproben-Methode nicht wahlen.

Hyperparasitism by virgin female Encarsia tricolor was studied by direct observation of its behaviour when contacting two secondary host species (Encarsia formosa and E. tricolor) at different host stages (first and second larval stage, third larval stage, and pupal stage). The searching and hyperparasitism behavioural sequence of E. tricolor was independent of the host stage of the whitefly (Aleyrodes proletella), and was similar to several related primary parasitoid species. In experiments with equal numbers of secondary hosts, encounter frequencies were equal for both secondary host species in all developmental stages observed. However, rates of hyperparastism were different according to host stage and host species. Hosts in the late larval stages were most preferred for hyperparasitization and the heterospecific E. formosa was more preferred as a secondary host than the conspecific, E. tricolor, in particular from the prepupal stage onwards. The window of vulnerability, i.e., the duration of the period in which a secondary host is susceptible to hyperparasitism, was largely determined by the occurrence and rate of melanization after the onset of pupation. The duration of a successful hyperparasitization event was longer than one that failed. Superparasitism occurred only once in all cases. The potential effect of autoparasitoids on biological control programs and the consequences for selection and release of an effective, yet ecologically safe agent are discussed.

Biological control is a valuable and effective strategy for controlling arthropod pests and has been used extensively against invasive arthropods. As one approach for control of invasives, exotic natural enemies from the native range of a pest are introduced to areas where control is needed. Classical biological control began to be used in the late 1800s and its use increased until, beginning in 1983, scientists began raising significant concerns and questions about nontarget and indirect effects that can be caused by these introductions. In recent years, similar issues have been raised about augmentative use of exotic natural enemies. Subsequently, international guidelines, national regulations and scientific methods being used for exotic natural enemies in biological control have changed to require appropriate specificity testing, risk assessment and regulatory oversight before exotic natural enemies can be released. National and international standards aimed at minimizing risk have increased awareness and promoted more careful consideration of the costs and benefits associated with biological control. The barriers to the implementation of classical and augmentative biological control with exotic natural enemies now are sometimes difficult and, as a consequence, the numbers of classical biological control programs and releases have decreased significantly. Based in part on this new, more careful approach, classical biological control programs more recently undertaken are increasingly aimed at controlling especially damaging invasive arthropod pests that otherwise cannot be controlled. We examine evidence for these revised procedures and regulations aimed at increasing success and minimizing risk. We also discuss limitations linked to the apparent paucity of post-introduction monitoring and inherent unpredictability of indirect effects.

Location, structure and histology of chemosensilla on the tip of the ovipositor of the parasitoid Leptopilina heterotoma are described based on SEM and TEM studies. Furthermore, we developed a method for recording extracellular action potentials from the gustatory neurons in response to host haemolymph. This method allowed us to record multi-unit recordings from a sensillum occurring singly on the unpaired ovipositor valve. The TEM study of the ovipositor tip revealed the presence of six dendrites, the electrophysiological recordings provided evidence for the activity of three or possibly four gustatory neurons in response to the complex stimulus offered, leaving other taste functions or a mechanoreceptor function open for the remaining neurons.

The solitary parasitoid Leptopilina heterotoma is one of the best studied organisms concerning the ecology, behaviour and physiology of host discrimination. Behavioural evidence shows that L. heterotoma uses its ovipositor to discriminate not only between parasitized and unparasitized Drosophila melanogaster larvae, but also to discriminate between hosts with different numbers of parasitoid eggs. The existing knowledge about how and when the parasitoid marks the host motivated us to unravel the chemosensory basis of host discrimination by L. heterotoma that allows it to choose the “best” host available. In this paper we report on electrophysiological recordings of multi-neural responses from the single taste sensillum on the tip of the unpaired ovipositor valve. We stimulated this sensillum with haemolymph of unparasitized, one-time-parasitized and two-times-parasitized Drosophila larvae. We demonstrate for the first time that quantitative characteristics of the neural responses to these haemolymph samples differed significantly, implying that host discrimination is encoded by taste receptor neurons in the multi-neuron coeloconic ovipositor sensillum. The activity of three of the six neurons present in the sensillum suffices for host discrimination and support the hypothesis that L. heterotoma females employ an ensemble code of parasitization status of the host.

Classical optimal-foraging theory predicts that a parasitoid is less likely to leave a patch after a host encounter when the host distribution is aggregated, whereas a parasitoid is more likely to leave after a host encounter when the host distribution is regular. Field data on host distributions in the area of origin of the whitefly parasitoid Encarsia formosa showed that whiteflies aggregate at several spatial scales. However, infested leaves most likely contained a single host. This suggests that a host encounter is not enough to decide when to leave. We therefore tested the effect of host distribution and parasitoid experience on patch-leaving behavior. Each parasitoid was observed for several consecutive days in a three-dimensional arena with leaflets containing on average one host per leaflet in an either regular or aggregated host distribution. A proportional hazards model showed that a host encounter decreased the leaving tendency on a leaflet with one host when the time since the latest host encounter was short, but increased the leaving tendency when the time since the latest host encounter was long, independent of host distribution. We conclude that a parasitoid can switch from decreasing to increasing its tendency to leave a patch after a host encounter. We propose two hypotheses that may explain the evolution of such a switching mechanism. [KEYWORDS: optimal foraging ; patch-leaving behavior ; host distribution ; Encarsia formosa ; whitefly]

Parasitoids as well as many of their herbivorous hosts, depend on carbohydrate-rich food during the adult stage. Different types of nectar and honeydew vary with regard to their sugar composition. In order to successfully exploit a food source, the insect must show a positive gustatory response to its component sugars and be able to digest and metabolise them. Here we tested the herbivore Plutella xylostella L. (Lepidoptera: Plutellidae) and its parasitoid Diadegma semiclausum (Hellén) (Ichneumonidae: Campopleginae) with respect to their feeding response and longevity when provided with one of nine sugars (fructose, glucose, lactose, maltose, melibiose, melezitose, raffinose, sucrose, or trehalose). Both species responded to and showed a prolonged life span on a broad range of sugars. The impact of food supply on life span was about sixfold higher for the parasitoid than for the herbivore. In general, there was a good fit between gustatory response and achieved longevity, with some outliers. Both species showed only weak responses to melibiose, which significantly prolonged life span. The parasitoid showed a gustatory response to melezitose, which did not prolong its life span. The parasitoid and its herbivorous host responded differently to trehalose. These differences in gustatory response and longevity show the potential for application of selective sugar sources in conservation biological control. At the same time it also reveals a risk that the indiscriminate application of sugar sources may stimulate herbivory. [KEYWORDS: herbivore ; selectivity ; food supply ; conservation biological control ; Lepidoptera ; Plutellidae ; Ichneumonidae ;Campopleginae]

Insect parasitoids lay their eggs in arthropods. Some parasitoid species not only use their arthropod host for oviposition but also for feeding. Host feeding provides nutrients to the adult female parasitoid. However, in many species, host feeding destroys an opportunity to oviposit. For parasitoids that attack Homoptera, honeydew is a nutrient-rich alternative that can be directly imbibed from the host anus without injuring the host. A recent study showed that feeding on host-derived honeydew can be an advantageous alternative in terms of egg quantity and longevity. Here we explore the conditions under which destructive host feeding can provide an advantage over feeding on honeydew. For 5 days, Encarsia formosa Gahan (Hymenoptera: Aphelinidae) parasitoids were allowed daily up to 3 h to oviposit until host feeding was attempted. Host feedings were either prevented or allowed and parasitoids had ad libitum access to honeydew between foraging bouts. Even in the presence of honeydew, parasitoids allowed to host feed laid more eggs per hour of foraging per host-feeding attempt than parasitoids that were prevented from host feeding. The higher egg-laying rate was not compromised by survival or by change in egg volume over time. In conclusion, host feeding can provide an advantage over feeding on honeydew. This applies most likely under conditions of high host density or low extrinsic mortality of adult parasitoids, when alternative food sources cannot supply enough nutrients to prevent egg limitation. We discuss how to integrate ecological and physiological studies on host-feeding behavior.

Conocer el comportamiento de búsqueda de sus hospederos por parte de los parasitoides es un paso esencial para estimar la eficacia de estos agentes de control biológico y poder desarrollar estrategias para su liberación en el cultivo. En el presente trabajo se estudió el comportamiento de búsqueda de ninfas de Trleteurodes vaporariorum por parte del parasitoide A. fuscipennis en plantas de frijol en condiciones de laboratorio. Se determinó el tiempo de residencia del parasitoide y su actividad de búsqueda (%) en plantas asperjadas y no asperjadas con un fungicida sulfurado e infestadas o no con ninfas de T. vaporariorum. El tiempo de permanencia del parasitoide y su actividad de búsqueda en las plantas infestadas fue mayor que en plantas no infestadas o asperjadas. El tiempo de permanencia del parasitoide se prolongó por dos eventos: encuentros con hospederos no parasitados y toma de alimento de la planta. Los resultados sugieren que: 1) El fungicida sulfurado perturbó al parasitoide haciendo que abandonara más rápidamente la planta. Si A. fuscipennis es utilizado en un programa de Manejo Integrado de Plagas (MIP) para mosca blanca, se debe tener cuidado de que su liberación o migración al cultivo no ocurra inmediatamente después de la aplicación de un fungicida sulfurado; 2) A. fuscipennis, a pesar de ser un parasitoide pro-ovigénico que nace eón su carga de huevos lista para ser ovipositada, necesita alimentarse de la planta hospedera, posiblemente para compensar el gasto energético ocasio• nado por can mar activamente en busca de sus hospederos.

Persistence of Aschersonia aleyrodis, a fungal pathogen of whitefly, was studied on cucumber, gerbera and poinsettia. Germination capacity and infectivity of conidia, which stayed on the plants for up to 1 month, were assessed. Average germination of conidia on the leaves was low (< 14%), whereas most of the conidia transferred from the leaf to water agar were viable, even after having been on the leaf surface for 1 month. Germination capacity was highest on cucumber, followed by poinsettia and lowest on gerbera. On cucumber leaves, conidia stayed viable and were able to infect 90% of whitefly nymphs 31 d after spore application. On gerbera, germination capacity decreased considerably from 80% (day 0) to 40% (day 31). This was reflected in nymphal mortality, which declined from 75% to 40%. Despite the high germination capacity (60%) of conidia on poinsettia after an exposure of one month, nymphal mortality decreased from 70% at the day of spore application to 10% after 3 d at leaf surface, and remained low throughout the monitoring period. Relations between germination capacity, infectivity and the host plant environment such as phyllosphere microorganisms, secondary plant metabolites and microclimate are discussed.

Observations on the host attack behaviour of the parasitoid Leptopilina heterotoma (Hymenoptera : Eucoilidae) led to the supposition that this wasp should possess a structure on its ovipositor by which it can hold a host larvae in a fixed position until the larva is paralyzed. The ovipositor was studied by light, scanning and transmission electron microscopy and appeared to have a clip with teeth on the unpaired valve of the ovipositor, about 50 μm from the tip. Based on the structure of this ovipositor clip and the oviposition behaviour, the functioning of the ovipositor clip is discussed.

Orius laevigatus (Hemiptera: Anthocoridae) is a key predator of thrips and is mass reared in large numbers for use in biological control. The aim of this study was to evaluate the effect of founder population size on the biological and behavioral performance of O. laevigatus over time. Laboratory lines were started from 1, 10 and 50 founder couples from 750 adults collected in the field and their performance was evaluated at the 5th–6th and 10th–11th generations. Adaptation to the captive rearing situation occurred in the 10 and 50 founder couples lines while it failed in the 1 founder couple line. The intrinsic rate of natural increase (rm) increased and the period for doubling the population (D) decreased over the generations in the 10 and 50 founder couples lines, while (rm) decreased and (D) increase in the 1 founder couple line. Also, consumption of Frankliniella occidentalis prey was significantly lower for females from the 1 founder couple line at the 5th generation compared to females from the 10 and 50 founder couples lines. Females of laboratory lines of all founder couples did not respond to odours from thrips infested plants during the 5th and 10th generations, whereas wild females strongly reacted to these odours. We suggest that the lack of reaction to infested plant volatiles may be due to the artificial rearing method where mass reared predators do not experience an infested crop. The results showed that the 1 founder couple line differed from the 10 and 50 founder couples lines, suggesting that bottlenecking had an effect at that level. However, no difference was found between the 10 and 50 founder couples lines which suggest that these founder numbers can be used to start laboratory-reared O. laevigatus lines without a significant loss in quality of its relevant biological characteristics.

In Europe, commercial use of biological control has seen a very fast development during the past 30 years. Currently, Europe has 26 companies which produce more than 80 species of natural enemies. The 29 most commonly sold natural enemies are discussed in this survey. The recommended release rate, the unit of sale, the target pest(s), and the number of companies producing a certain natural enemy are specified. Large differences in prices for biological control agents exist between Europe and the United States. Many more species of natural enemies are commercially available in Europe than in the United States, mainly as a result of the much larger greenhouse industry in Europe.

We tested the hypothesis that two populations of the parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae) differed in their behavioral interactions with the whitefly host Bemisia tabaci Gennadius (Homoptera: Aleyrodidae). The two wasp populations were studied because previous work suggested large differences between the populations in performance on this host. In this study the populations differed behaviorally in both the number of hosts encountered and their reactions to hosts once encountered. The population reared for many years on Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae) that previously performed more poorly on B. tabaci had a higher host encounter rate but rejected hosts more frequently. The population reared for a number of years on B. tabaci encountered fewer hosts but accepted a higher percentage of hosts for oviposition. The number of parasitized hosts did not differ between the two populations, however. These data demonstrate that there are heritable differences between these two populations of asexual wasps in host-associated behavioral traits. These behavioral differences in host acceptance do not explain performance differences seen in the earlier study, however, possibly due to different conditions between the two experiments.

International audience; With the establishment of the Plant Health Panel in 2006, EFSA became the body responsible for risk assessment in the plant health area for the European Union (EU). Since then more than 70 outputs have been produced dealing with the full range of organisms harmful to plant health across all crop types and plants in the environment. There has been an increasing trend towards producing scientific opinions which are full pest risk assessments for the whole EU territory. In its work, and as a contribution to the wider development of risk assessment methodology, the Panel has developed a series of guidance documents. These deal with the peer review of existing pest risk assessments, a framework for conducting risk assessments which harmonise standards set by the International Plant Protection Convention and the legislative requirements of the EU, and extension of this framework to include environmental risk assessment and the evaluation of risk reducing options. Quantitative approaches have become increasingly important during this time. The Panel has developed such methods in climatic mapping (in association with the Joint Research Councils), application of spatial spread models, re-evaluation of quantitative pathway analyses, and in statistical modelling of experimental data. A Plant Health Network has been established to facilitate interaction with EU Member States, especially in relation to data collection and co-ordination of risk assessment activities. At the current time a revision of the EU Plant Health Regime is being formulated. The legislative consequences of the revision will be of considerable significance for the work of the Plant Health Panel.

Since the establishment of Harmonia axyridis in Europe, populations of native ladybirds have decreased. Overwintering survival is one of the aspects of the biology of H. axyridis that may contribute to its firm establishment in and invasion of a new area. In this study winter survival of five wild H. axyridis populations was assessed under natural and semi-natural conditions, with a focus on the potential influence of location and orientation on winter survival. Overwintering survival of H. axyridis in the Netherlands is high: 70.8–88.2%. When overwintering at one central site, populations sampled at five locations showed statistically significant different mortality rates. Furthermore, winter survival of H. axyridis at the sample sites was higher when beetles were hibernating at the southwestern sides of buildings, where most aggregations of ladybirds were found. Survival was higher at sheltered sites compared to exposed sites. Harmonia axyridis has a comparable or higher overwintering survival than most common native ladybird species. A high overwintering survival results in a large post-hibernation population in spring, leading to a rapid population build-up. Thus, the high winter survival probably contributes to the success of the exotic H. axyridis .

Harmonia axyridis has been introduced as a biological control agent in Europe and the USA. Since its introduction, it has established and spread, and it is now regarded as an invasive alien species. It has been suggested that intraguild predation is especially important for the invasion success of H. axyridis. The aim of this study was to compare the intraguild predation behaviour of three ladybird species (Coccinella septempunctata, Adalia bipunctata, and H. axyridis). Predation behaviour was investigated in semi-field experiments on small lime trees (Tilia platyphyllos). Two fourth-instar larvae placed on a tree rarely made contact during 3-hour observations. When placed together on a single leaf in 23%–43% of the observations at least one contact was made. Of those contacts 0%–27% resulted in an attack. Harmonia axyridis attacked mostly heterospecifics, while A. bipunctata and C. septempunctata attacked heterospecifics as often as conspecifics. In comparison with A. bipunctata and C. septempunctata, H. axyridis was the most successful intraguild predator as it won 86% and 44% of heterospecific battles against A. bipunctata and C. septempunctata respectively, whilst A. bipunctata won none of the heterospecific battles and C. septempunctata won only the heterospecific battles against A. bipunctata. Coccinella septempunctata dropped from a leaf earlier and more often than the other two species but was in some cases able to return to the tree, especially under cloudy conditions. The frequency with which a species dropped did not depend on the species the larva was paired with. The results of these semi-field experiments confirm that H. axyridis is a strong intraguild predator as a consequence of its aggressiveness and good defence against predation from heterospecific species. The fact that H. axyridis is such a strong intraguild predator helps to explain its successful establishment as invasive alien species in Europe and the USA.

The effectiveness of insecticide-treated bed nets and indoor insecticide sprays to control adult mosquito vectors is being threatened by the spread of insecticide resistance. We argue for expanding beyond “insecticide monotherapy” to more sustainable integrated vector management strategies that use optimal suites of control tactics. Experience in agriculture suggests that such integrated approaches can provide more effective and durable pest management. This shift will require increased investment in research and translational science. Failure to act risks a resurgence of malaria and erosion of community support and donor commitment.

The interaction between the entomopathogenic fungus Aschersonia aleyrodis and the parasitoid Encarsia formosa on greenhouse whitefly as a host organism was studied, in particular, the survival of the parasitoid after treatment of parasitized hosts with fungal spores. The mean number of parasitized black pupae per parasitoid produced at 25°C was significantly reduced after spore treatment in the first three days following parasitization. Spore treatment four, seven or ten days after parasitization resulted in a mean number of parasitized pupae not significantly different from the number of black pupae in the control. The rather sudden change from low to high survival of parasitized hosts when treated with spores four days after parasitization in spite of high numbers of infected unparasitized larvae, coincided with the hatching of the parasitoid larva from the egg inside the host. Possible reasons for this decrease in susceptibility to infection after parasitoid egg hatch, such as induced changes in host cuticle or haemolymph, are discussed. Parasitoids emerged from treated hosts did not show differences in reproduction compared with parasitoids emerging from untreated hosts. Both natural enemies of whitefly are compatible to a great extent.