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Tuta absoluta (Meyrick), a key pest of tomato, is quickly spreading over the world and biological control is considered as one of the control options. Worldwide more than 160 species of natural enemies are associated with this pest, and an important challenge is to quickly find an effective biocontrol agent from this pool of candidate species. Evaluation criteria for control agents are presented, with the advantages they offer for separating potentially useful natural enemies from less promising ones. Next, an aggregate parameter for ranking agents is proposed: the pest kill rate km. We explain why the predator's intrinsic rate of increase cannot be used for comparing the control potential of predators or parasitoids, while km can be used to compare both types of natural enemies. As an example, kill rates for males, females and both sexes combined of three Neotropical mirid species (Campyloneuropsis infumatus (Carvalho), Engytatus varians (Distant) and Macrolophus basicornis (Stål)) were determined, taking all life-history data (developmental times, survival rates, total nymphal and adult predation, sex ratios and adult lifespan) into account. Based on the value for the intrinsic rate of increase (rm) for T. absoluta and for the kill rate km of the predators, we predict that all three predators are potentially able to control the pest, because their km values are all higher than the rm of the pest. Using only km values, we conclude that E. varians is the best candidate for control of T. absoluta on tomato, with C. infumatus ranking second and M. basicornis last.

Biological control activities on 18 (groups of) Caribbean islands are summarized. Many natural enemies were introduced to these islands through Trinidad and Tobago up to 1980. Also, inter-island exchange of biocontrol agents took place. The majority of projects concerned classical biocontrol, while in some cases natural, conservation and augmentation biocontrol were used. Successes were obtained with biocontrol of pests in crops such as arrowroot, citrus, coconut, cotton and sugarcane and of weeds like prickly pear and puncture vine. After 1980, the number of natural enemy introductions decreased, though the region was faced with many invasions by exotic pests, including the citrus leaf miner, citrus blackfly, papaya mealybug, giant African snail, coconut whitefly and pink hibiscus mealybug. Two large region-wide programmes resulted in successful biocontrol of the pink hibiscus mealybug and the papaya mealybug. In addition, biocontrol by a native natural enemy complex was demonstrated for the coconut whitefly and the passion vine mealybug. The Food and Agriculture Organization of the United Nations (FAO) Code of Conduct for the Import and Release of Exotic Biological Control Agents has recently been applied in the region. Farmers Field Schools, with the aim to enable farmers to use IPM and become less dependent on chemical pesticides, are being implemented.

A number of introductions of parasitoids and predators were carried out in the 1950s for classical biological control of olive scale, woolly apple aphid, white peach scale, Mediterranean fruit fly and Anastrepha fruit fly, with control of cottony cushion scale by the coccinellid Rodolia being a particular success. In 1963, dipteran natural enemies were introduced for the control of sugarcane borers, Diatraea spp. Since 1963 native hymenopteran and dipteran parasitoids have been field collected and re-released for control of the borers. In 1969 an IPM programme of sugarcane borers was started, and biocontrol in sugarcane in the period 1970-2000 mainly consisted of augmentative releases of hymenopteran and tachinid parasitoids. Another successful IPM programme dealt with control of potato moth species with a product - now commercially available - that contains a native strain of the granulosis virus Baculovirus phthorimaea and a native strain of Bacillus thuringiensis. Coffee berry borer was brought under biocontrol in the 1990s by releasing a hymenopteran parasitoid and application of an entomopathogenic fungus. An increased demand for organic products since 2000 has stimulated work on isolation, characterization, mass production, formulation and certification of a number of microbial control agents. These are used in many crops and examples are microbial control of pest in potato and quinoa. Many of the quinoa pests are kept under natural control by predators and parasitoids, which has been well documented during the past 10 years. Currently most pests in sugarcane and soybean are under a combination of natural, augmentative and classical biocontrol.

The egg is the first life stage directly exposed to the environment in oviparous animals, including many vertebrates and most arthropods. Eggs are vulnerable and prone to mortality risks. In arthropods, one of the most common egg mortality factors is attack from parasitoids. Yet, parasitoids that attack the egg stage are absent in more than half of all insect (sub)orders. In this review, we explore possible causes explaining why eggs of some insect taxa are not parasitized. Many insect (sub)orders that are not attacked by egg parasitoids lack herbivorous species, with some notable exceptions. Factors we consider to have led to escape from egg parasitism are parental egg care, rapid egg development, small egg size, hiding eggs, by e.g. placing them into the soil, applying egg coatings or having thick chorions preventing egg penetration, eusociality, and egg cannibalism. A quantitative network analysis of host–parasitoid associations shows that the five most-speciose genera of egg parasitoids display patterns of specificity with respect to certain insect orders, especially Lepidoptera and Hemiptera, largely including herbivorous species that deposit their eggs on plants. Finally, we discuss the many counteradaptations that particularly herbivorous species have developed to lower the risk of attack by egg parasitoids.

In the 1960s, natural control was shown to play an important role in the reduction of saturniid populations in Spondias fruit trees by dipteran and hymenopteran parasitoids and in the reduction of coconut weevil populations by a staphelinid. Later, other cases of high degrees of natural control were documented, such as that of the cotton leafworm by a native egg parasitoid and of the Mexican beetle by an ectoparasitic mite as well as by a pupal parasitoid. Classical biocontrol of citrus blackfly and purple scale in citrus was initiated in the 1970s, as well as augmentative biocontrol of lepidopteran pests with egg parasitoids. Another, very successful, augmentative biocontrol project during the 1970s concerned the killing of two mosquito species by an entomopathogenic nematode. Prospecting for nematophagous fungi in 1977 in El Salvador resulted in finding 18 species. Recently, several microbial control agents have been produced by the Escuela Agrícola Panamericana, Zamorano in Honduras for control of soil-borne diseases; nematodes and arthropod pests have been registered and are applied in El Salvador. Tilapia fish are used for biocontrol of Aedes mosquitoes, the vector of, among others, Zika, dengue and chikungunya viral diseases.

Biological control was initiated in Guatemala in the 1990s, after costs for chemicals had increased substantially and had resulted in a drastic decrease in production of, for example, cotton and tomato. Most often, augmentative biocontrol is used in Guatemala. Examples of successful programmes are control of: (i) lepidopterans in cotton and vegetables by microbial agents and egg parasitoids; (ii) cotton leafworm by egg parasitoids and a predator in cotton; (iii) coffee berry borer with a parasitoid and a microbial agent; (iv) sugarcane borer with a parasitoid and a microbial agent; (v) diamondback moth with parasitoids in cruciferous crops; and (vi) white grub in maize with an entomopathogenic nematode. In 1998, populations of Anopheles mosquitoes, which are vectors of malaria, were successfully reduced by killing their larvae with microbial agents, resulting in a 50% lower malaria prevalence. A recently started project concerns classical biocontrol of the Asian citrus psyllid with a parasitoid. Biocontrol is expected to grow because of, inter alia, demands for residue-free food by countries to which Guatemala exports.

This volume summarises current developments in integrated pest management (IPM), focussing on insect pests. Chapters discuss advances in understanding species and landscape ecology on which IPM is founded. The book then reviews advances in cultural, physical and, in particular, biological methods of control. Topics include developments in classical, conservation and augmentative biological control as well as the use of entomopathogenic fungi, viruses, nematodes and semiochemicals. The final parts of the book summarise current research on monitoring pesticide use as well as emerging classes of biopesticides.

The Nagoya Protocol is a supplementary agreement to the Convention on Biological Diversity that provides a framework for the effective implementation of the fair and equitable sharing of benefits arising out of the utilization of genetic resources, including invertebrate biological control agents. The Protocol came into force on 12 October 2014, and requires signatories and countries acceding to the Protocol to develop a legal framework to ensure access to genetic resources, benefit-sharing and compliance. The biological control community of practice needs to comply with access and benefit sharing regulations arising under the Protocol. The IOBC Global Commission on Biological Control and Access and Benefit Sharing has prepared this best practices guide for the use and exchange of invertebrate biological control genetic resources for the biological control community of practice to demonstrate due diligence in responding to access and benefit sharing requirements, and to reassure the international community that biological control is a very successful and environmentally safe pest management method based on the use of biological diversity. We propose that components of best practice include: collaborations to facilitate information exchange about what invertebrate biological control agents are available and where they may be obtained; knowledge sharing through freely available databases that document successes (and failures); cooperative research to develop capacity in source countries; and transfer of production technology to provide opportunities for small-scale economic activity. We also provide a model concept agreement that can be used for scientific research and non-commercial release into nature where access and benefit sharing regulations exist, and a model policy for provision of invertebrate biological control agents to other parties where access and benefit sharing regulations are not restrictive or do not exist.

The predators Macrolophus basicornis (Stal), Engytatus varians (Distant) and Campyloneuropsis infumatus (Carvalho) consume large numbers of tomato pests such as Bemisia tabaci (Gennadius) and Tuta absoluta (Meyrick). However, they are zoophytophagous and feed on plant parts as well. We evaluated the type and effect of injury caused by nymphs and adults of these mirids on tomato seedlings and fruit in the absence of prey. For each mirid species, seedlings were exposed to groups of 20 nymphs or adults for 72 h, and fruits were exposed for 48 h to groups of four nymphs or adults. Type and the number of injury on stems, petioles and leaflets of tomato seedlings and fruits were recorded after removal of insects. Nymphs and adults of these mirids caused necrotic rings on the leaflets, but no injury was observed on stem and petioles. The necrotic rings on leaflets consisted of blemishes, characterized by feeding punctures surrounded by a yellowish, bleached area. The number of necrotic rings did not exceed one per individual mirid and seedlings developed normally. Nymphs also caused feeding punctures on tomato fruit, but in even lower numbers than on leaflets. Two weeks after the start of the experiment the tomato fruit still looked fresh and feeding punctures had disappeared. Adults did not cause any injury to tomato fruit. The results indicate that nymphs and adults of these zoophytophagous mirids cause little injury to tomato seedlings and fruit, even when present in high densities and in the absence of prey, making them interesting candidates for biological control.

The Asian ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is regarded as an invasive species in many parts of the world. In a previous study we hypothesised that H. axyridis enters diapause at the end of October and then shifts to a quiescent state in December in northwestern Europe. In the present study we test this idea of a short, early period of diapause by sampling beetles from their hibernation sites immediately after their migratory flights in October, subsequently keeping them in outdoor cages, and then, after certain time-intervals, measuring the pre-oviposition time under optimal egg-laying laboratory conditions at 25 degrees C. We did this at both short (12L) and long (16L) photoperiods, since a photoperiodic response is an indicator of true diapause, rather than quiescence. A significant, albeit small, difference in pre-oviposition period between the two photoperiods, which disappears in December, corroborates our earlier hypothesis that the ladybirds are in a state of diapause until mid-December. Compared with that of native ladybirds the diapause of H. axyridis generally is relatively short and weak; moreover, it appears to have become shorter over the last decade. This flexibility in diapausing behaviour may be an important factor that contributes to the invasive success of H. axyridis.

Behaviour of nocturnal insects is routinely observed under red light, but it is unclear how the behaviour under red light compares to behaviour in complete darkness, or under a source of white light. Here, we measure movement behaviour of the nocturnal carabid beetle Pterostichus melanarius Illiger (Coleoptera: Carabidae) using camera recording under a near-infrared (nir), red or white radiation source. Red light significantly reduced movement speed in females similar to the effect of white light and different from nir. Also movement activity and pause length were affected by radiation source, with a significant difference between nir and white light, and with intermediate values in red light. The results presented here indicate that P. melanarius has different movement behaviour under the three radiation sources and suggest that nir rather than red radiation is most appropriate for measuring behaviour in total darkness. However, in the field total darkness is rare both because of natural light sources such as the moon and stars but increasingly also because of ecological light pollution, and therefore red light may still be of use for observing ecologically and practically relevant natural night-time behaviour.

Correct identification of the source population of an invasive species is a prerequisite for testing hypotheses concerning the factors responsible for biological invasions. The native area of invasive species may be large, poorly known and/or genetically structured. Because the actual source population may not have been sampled, studies based on molecular markers may generate incorrect conclusions about the origin of introduced populations. In this study, we characterized the genetic structure of the invasive ladybird Harmonia axyridis in its native area using various population genetic statistics and methods. We found that native area of H. axyridis most probably consisted of two geographically distinct genetic clusters located in eastern and western Asia. We then performed approximate Bayesian computation (ABC) analyses on controlled simulated microsatellite data sets to evaluate (i) the risk of selecting incorrect introduction scenarios, including admixture between sources, when the populations of the native area are genetically structured and sampling is incomplete and (ii) the ability of ABC analysis to minimize such risks by explicitly including unsampled populations in the scenarios compared. Finally, we performed additional ABC analyses on real microsatellite data sets to retrace the origin of biocontrol and invasive populations of H. axyridis, taking into account the possibility that the structured native area may have been incompletely sampled. We found that the invasive population in eastern North America, which has served as the bridgehead for worldwide invasion by H. axyridis, was probably formed by an admixture between the eastern and western native clusters. This admixture may have facilitated adaptation of the bridgehead population.

The effect of temperature on the biology of Praon volucre (Haliday, 1833) (Hymenoptera: Braconidae) in Macrosiphum euphorbiae (Thomas, 1878) (Hemiptera: Aphididae) hosts was studied and the thermal requirements of the parasitoid were determined. Experiments were carried out at 16, 19, 22, 25, and 28 ± 1°C with 70 ± 10% relative humidity and 12 h photophase. The developmental time of P. volucre males and females decreased between 16 and 23°C. The highest percentages of aphid parasitism and adult parasitoid emergence were observed at 16, 19, and 22°C. The sex ratio (expressed as % females) decreased between 16 and 23°C. Male and female longevity was high between 19 and 22°C and decreased strongly at 25°C. The lower temperature threshold for P. volucre was 5.17°C and the thermal constant was 243 degrees-days.

This paper briefly describes the foundations and characteristics of biological control in protected cropping and what drivers are behind adoption of this management system within this industry. Examining a brief history of biological control in greenhouses and what makes it a successful management strategy within the industry, the authors describe the rapid growth of biological control in parts of Europe and what this may mean for the industry in other parts of the world. The reaction of the greenhouse industry to several consumer led campaigns aimed at reducing the incidence of pesticides in the marketplace may be replicated in many other parts of the world. The size and robustness of the biological control industry in greenhouses, which is a reflection of the inherent characteristics of this industry that lends itself to biological control, is strong and growing with indications that this trend will be followed in many areas of the world.

Zusammenfassung Zur Parasit-Wirt-Beziehung zwischen Encarsia formosa (Hym., Aphelinidae) und Trialeurodes vaporariorum (Hom., Aleyrodidae). XII. Populationsdynamik des Parasiten und des Wirtes in einem grosen Gewachshaus, unduberprufung der in den Niederlandenublichen Aussetzungsmethoden des Parasiten Die weise Fliege, Trialeurodes vaporariorum (Westwood), ist ein Grosschadling an verschiedenen Gewachshauskulturen. Seit etwa 1970 wird dieser Schadling auf wachsender Flache und in immer mehr Landern durch Einsatz eines naturlichen Feindes, der Schlupfwespe Encarsia formosa Gahan bekampft. 1979 wurde dieses biologische Verfahren in 11 Landern auf ungefahr 1200 ha durchgefuhrt. Einige Jahre nach Beginn der Anwendung von E. formosa begannen wir eine Forschungsarbeituber den Massenwechsel des Parasiten und seines Wirtes, mit folgenden Zielsetzungen: 1 Entwicklung einer schnellen und zuverlassigen Zahlmethode fur Gartner sowie Lieferanten der Schlupfwespen. 2 Bewertung der in den Niederlandenublichen Methode des Schlupfwespeneinsatzes. 3 Erklarung der gewohnlich unbefriedigenden Erfolge in kleineren Gewachshausern im Vergleich zu Groseinheiten. 4 Bewertung der Hypothese von Huffaker (1958), wonach die Co-Existenz von Beutetier und Jager nur moglich ware, wenn bestimmte Bedingungen bezuglich raumlicher Verteilung und Wanderfahigkeit von beiden gegeben sind. Der Massenwechsel und die Verteilung der Weisen Fliege und ihres Parasiten wurde 16 Wochen lang in einem 6480 qm grosen Gewachshaus mit 18 000 Tomatenpflanzen verfolgt. Dabei wurden 2 Verfahren verglichen: die wochentliche Zahlung der Puppen an allen befallenen Pflanzen, und eine geschichtete Stichprobe mit wochentlicher Kontrolle an 0,5% der Pflanzen. Da das letztere Verfahren die wirklichen Zahlen und Verteilungen der beiden Insekten nicht erfaste, darf es den Praktikern nicht empfohlen werden (Eggenkamp-Rotteveel Mansveld et al. 1978). Die Zahlungen und die daraus konstruierten Verteilungsmuster ermoglichten folgende Schlusfolgerungen: 1 Das in den Niederlandenubliche Einsatzschema war erfolgreich. Angefangen wird dabei 1 bis 4 Wochen nach dem ersten Auftreten des Schadlings; insgesamt werden im allgemeinen 6 Schlupfwespen pro Pflanze eingesetzt, verteiltuber 4 Freilassungen in 14tagigem Abstand; in unserem Fall waren es jedoch insgesamt 9,8 Schlupfwespen. Die (schwankende) Zahl der Weisen Fliegen blieb sehr gering unduberschritt niemals 0,1 Puppe pro Pflanze. Bis zum Ende der Kultur wurde kein von der Weisen Fliege verursachter Schaden beobachtet. Ein solcher Erfolg ist keine experimentelle Ausnahme, sondern durchaus in der Praxis erzielbar. 2 Sowohl die Weisen Fliegen als auch die Schlupfwespen zeigten eine ausgepragte Herdbildung. Die Herde veranderten sich allmahlich zu einem regelmasigen Verteilungsmuster in der letzten Woche. Dies wurde ausgelost von den wandernden Imagines der Weisen Fliege. Dadurch wuchs die Zahl der besiedelten Pflanzen; weil aber zugleich die Zahl der Puppen pro besiedelte Pflanze abnahm, stieg die Gesamtzahl der Weisen Fliegen nicht an. 3 Die aktive Verbreitung und die Suchfahigkeit der Schlupfwespen ist ausgezeichnet. Das einzelne Insektuberfliegt Abstande von mindestens 10 Metern. Sogar vereinzelte Puppen des Wirtes innerhalb von Gruppen sonst unbesiedelter Pflanzen wurden aufgefunden. 4 Bei Huffakers (1958) Versuchen wurden Subpopulationen einer phytophagen Milbe manchmal vollstandig von einer rauberischen Milbe vernichtet, wahrend zu gleicher Zeit neue Subpopulationen des Phytophagen an anderen Fruchten hochkamen. Die Herde in unserem Versuch (Herd = Gruppe von Weisen Fliegen und/oder Schlupfwespen, die durch unbesiedelte Pflanzen wenigstens 3 m von anderen Gruppen getrennt ist) wurden nie ganzlich beseitigt. Innerhalb eines Herdes aber verschwand eine Subpopulation an einer Pflanze manchmal durch vollstandigen Parasitismus, wahrend sich an einer anderen Pflanze eine neue Subpopulation entwickelte. Von Herden oder Pflanzen, an denen alle Puppen parasitiert sind, wandern die Schlupfwespen ab und suchen sich neue Subpopulationen der Weisen Fliege. Dadurch schwankte die Parasitierungsrate pro Pflanze zwischen 0 und 100%, doch blieb die durchschnittliche Rate recht konstant (etwa 50%). Obwohl die Ereignisse an einzelnen Pflanzen sich also oft drastisch voneinander unterschieden, addierten die lokalen Unterschiede sich zu einer stabilen Lage im gesamten Gewachshaus.

The heteropteran predator Geocoris punctipes (Say) has been used in augmentative biological control since 2000 to control Lepidoptera. However, surprisingly, few data are available about the influence of temperature on its population development, which is of key importance to plan the number and moment of releases to obtain sufficient pest reduction. The objective of this study was to evaluate daily and total fecundity, longevity and life table parameters (mx, lx, rm, R, λ, T and TD) of G. punctipes at constant (16.8°C, 21.5°C, 24.5°C and 28.3°C) and corresponding varying (day/night) (21/11°C, 24/18°C, 27/21°C and 30/26°C) temperatures. Pairs of adult predators aged 24 h and originating from nymphs exposed to the same temperature regimes were kept at the above-mentioned temperature regimes in Petri dishes containing Anagasta kuehniella (Zeller) eggs and an oviposition substrate. Tests were conducted in climatic chambers at the different temperature regimes and a RH 70 ± 10% and a 14L: 10D photoperiod. Reproduction, longevity and life table parameters were significantly affected by temperature, with clear differences between treatments at low (16.8°C, 21/11°C, 21.5°C, 24/18°C) or a high (24.5°C, 27/21°C, 28.3°C, 30/26°C) temperature regimes. Highest reproduction and fastest population growth of G. punctipes took place at average temperatures ranging from 24.5°C to 30°C, and neither reproduction nor population growth was negatively influenced by varying temperatures at any of the temperature regimes.

Quantification of the movement of insects at field and landscape levels helps us to understand their ecology and ecological functions. We conducted a meta-analysis on movement of carabid beetles (Coleoptera: Carabidae), to identify key factors affecting movement and population redistribution. We characterize the rate of redistribution using motility μ (L2 T−1), which is a measure for diffusion of a population in space and time that is consistent with ecological diffusion theory and which can be used for upscaling short-term data to longer time frames. Formulas are provided to calculate motility from literature data on movement distances. A field experiment was conducted to measure the redistribution of mass-released carabid, Pterostichus melanarius in a crop field, and derive motility by fitting a Fokker–Planck diffusion model using inverse modelling. Bias in estimates of motility from literature data is elucidated using the data from the field experiment as a case study. The meta-analysis showed that motility is 5.6 times as high in farmland as in woody habitat. Species associated with forested habitats had greater motility than species associated with open field habitats, both in arable land and woody habitat. The meta-analysis did not identify consistent differences in motility at the species level, or between clusters of larger and smaller beetles. The results presented here provide a basis for calculating time-varying distribution patterns of carabids in farmland and woody habitat. The formulas for calculating motility can be used for other taxa.

Host-handling behavior is an important aspect of parasitoid foraging behavior. When a parasitoid encounters a potential host, the handling behavior starts with the evaluation of the host and continues if the host has been judged acceptable. Host handling is usually terminated after egg laying or host feeding and host marking. Host-handling behavior of an arrhenotokous population of two Eretmocerus species, E. mundus Mercet and E. eremicus Rose and Zolnerowich, along with a thelytokous population of E. mundus were compared under laboratory conditions. Several elements of host-handling behavior, including encountering, ascending, turning on host, descending, preening, egg laying, and host feeding were recorded. There were no correlations among the durations of these phases across parasitoid populations/species or host nymphal instars. Duration of different phases of host-handling behavior showed only slight and sometimes significant differences between different Eretmoceruspopulations/species. The actual laying of the egg had the longest duration of all host-handling behaviors, and was longer on third nymphal instars than on younger ones. Females of the three populations/species accepted the first three nymphal stages either for egg laying or for host feeding. Females spent a lot of time to make wounds in the host when preparing for host feeding, and eventually killed the host. The implications of these findings for the use of the different Eretmoceruspopulations/species in biological control are discussed.

Bemisia tabaci (Gennadius) is a serious pest of vegetable, ornamental, and agronomic crops throughout the world. To control B. tabaci, Eretmocerus eremicus Rose & Zolnerowich, and Eretmocerus mundus Mercet are considered the most effective parasitoids in dry tropical regions. In parasitoids, choosing the `right` hosts has direct consequences for their reproductive success and efficiency as biocontrol agent. Therefore, being able to discriminate a parasitized host from an unparasitized one would be important to prevent wasting time, eggs, and to reduce the mortality risk for their offspring. We evaluated intra- and interspecific host discrimination and the chance of super-parasitism or multi-parasitism in two populations of E. mundus (sexual and asexual) and E. eremicus. Different combinations and sequences of female introduction were carried out for the various populations and species. Experienced females avoided super-parasitism. However, naïve females did lay eggs under hosts that were previously parasitized by conspecific females. E. eremicus females avoided to multi-parasitize hosts parasitized by E. mundus. However, E. mundus females did multi-parasitize the hosts that had been parasitized earlier by E. eremicus. In the case of super-parasitism, the outcome showed that neither of the E. mundus populations was stronger, whereas in the case of multi-parasitism E. mundus appeared stronger than E. eremicus. Since those populations and species are morphologically similar a molecular method had to be developed to identify the outcome of super- or multi-parasitism, which is presented in Appendix A.

The authors most frequently credited for the European discovery of the parasitoid life cycle are Antoni van Leeuwenhoek, John Ray and Antonio Vallisnieri around the year 1700. Many other European authors published works on entomology in the 17th century and mentioned insects that we now recognize as parasitoids. Most of them were supposed until recently not to have understood the parasitoid life cycle. After rereading much of the old literature, we suggest this supposition is correct for, among others, Aldrovandi, Goedaert, Malphighi, and Redi. However, Lister, Merian, and Swammerdam (with the help of the painter Marsilius) all arrived at the correct interpretation of insect parasitism after observing most or all life history stages. The first correct interpretation of parasitism that we can trace, but which does not include the critical observation of oviposition by the adult female, is that of Swammerdam in 1669. The first recorded observation of oviposition that we can find is by the painter Marsilius but described by Swammerdam in 1678. However, this observation was not published until 1737-1738 by which time van Leeuwenhoek's influential 1700-1701 description had rendered it obsolete. Provisionally, we thus suggest Jan Jacob Swammerdam (assisted by Otto Marsilius) should be credited with the description of the discovery of the parasitoid life cycle in Europe. © 2004 Elsevier Inc. All rights reserved.

Differences in allelochemistry of plants may influence their ability to attract parasitoids.We studied responses of Diadegma semiclausum (Hellén), a parasitoid of the diamondback moth (Plutella xylostella L.), to inter- and intraspecific variation in odor blends of crucifers and a non-crucifer species. Uninfested Brussels sprout (Brassica oleracea L. gemmifera), white mustard (Sinapis alba L.), a feral Brassica oleracea, and malting barley (Hordeum vulgare L.) were compared for their attractivity to D. semiclausum in a Y-tube bioassay. Odors from all plants were more attractive to the parasitoid than clean air. However, tested against each other, parasitoids preferred the volatile blend from the three cruciferous species over that of malting barley.Wasps also discriminated between uninfested crucifers: mustard was as attractive as feral B. oleracea, and both were more attractive than Brussels sprout. Attractivity of uninfested plants was compared with that of plants infested by larvae of the host P. xylostella. Host-infested mustard and Brussels sprout were more attractive than uninfested conspecifics. Interestingly, the volatile blends of uninfested white mustard and infested Brussels sprout were equally attractive.We also compared the volatile composition of different plant sources by collecting headspace samples and analysing them with GC-MS. Similarities of volatile profiles were determined by hierarchic clustering and non-metric scaling based on the Horn-index. Due to the absence of several compounds in its blend, the volatile profile of barley showed dissimilarities from blends of crucifers. The odor profile of white mustard was distinctly different from the two Brassicaceae.Feral Brassica oleracea odor profile was different from infested Brussels sprout, but showed overlap with uninfested Brussels sprout. Odor blends from infested and uninfested Brussels sprout were similar, and mainly quantitative differences were found. D. semiclausum appears to discriminate based on subtle differences in volatile composition of odor blends from infested and uninfested plants. [KEYWORDS: Diadegma semiclausum ; Plutella xylostella ; Hordeum vulgare ; Sinapis alba ; Brassica oleracea ; olfactometer ; headspace volatiles ; GC-MS]

Life-history parameters (juvenile development time, adult longevity, host instar preference and rate of parasitism) offour parasitoids of Bemisia argentifolii (two strains of Encarsia formosa (D and B), Eretmocerus eremicus and Eretmocerus mundus) Were Studied in the laboratory. At 15degreesC juvenile development time was the shortest for E. formosa B (48 days), longest for E. eremicits (79.3 days) and intermediate for E. formosa D (62.8 days) and E. mundus (64 days) at 15degreesC. With increase in temperature, development time decreased to around 14 days for all species/strains at 32degreesC. The lower developmental threshold for development was 11.5, 8.1, 13.0 and 11.5degreesC for E. formosa D, E. formosa B, E. eremicus and E. mundus, respectively. E. formosa D and B, and E. mundus all appeared to prefer to parasitize 3(rd) instar nymphs. The presence of hosts shortened adult longevity in most of the parasitoids, with the exception of E. formosa B, which lived longer than other species/strains irrespective of the presence of hosts. At 15degreesC daily parasitism was very low by all parasitoids. The two Encarsia strains had a constant, but low rate of reproduction during adult life, while the two Eretmocerus species had a very high rate of reproduction when one-day old, which then decreased very quickly. Lifetime fecundity, estimated using a non-linear model, indicated that it was higher for the two Encarsia strains than for the Eretmocerus species. Life history parameters reported in the literature for the four parasitoids are reviewed and compared with Our results. Finally, the potential value for the biological control of whiteflies on greenhouse crops of parasitoids having either a high reproductive rate over a short period (Eretmocerus spp.) or a low rate of reproduction over a long period (Encarsia spp.) is discussed.

In the past 100 years many exotic naturalenemies have been imported, mass reared andreleased as biological control agents. Negativeenvironmental effects of these releases haverarely been reported. The current popularity ofinundative biological control may, however,result in problems, as an increasing number ofactivities will be executed by persons nottrained in identification, evaluation andrelease of biological control agents.Therefore, a methodology for risk assessmenthas been developed within the EU-financedproject `Evaluating Environmental Risks ofBiological Control Introductions into Europe[ERBIC]' as a basis for regulation of importand release of exotic natural enemies used ininundative forms of biological control (i.e.not in `classical biological control' thoughsome of the same principles and approachesapply). This paper proposes a general frameworkof a risk assessment methodology for biologicalcontrol agents, integrating information on thepotential of an agent to establish, itsabilities to disperse, its host range, and itsdirect and indirect effects on non-targets. Ofthese parameters, estimating indirect effectson non-targets will be most difficult, asmyriads of indirect effects may occur whengeneralist natural enemies are introduced. Theparameter `host range' forms a central elementin the whole risk evaluation process, becauselack of host specificity might lead tounacceptable risk if the agent establishes anddisperses widely, whereas, in contrast, amonophagous biological control agent is notexpected to create serious risk even when itestablishes and disperses well. Drawing onpublished information and expert opinion, theproposed risk assessment methodology is appliedto a number of biological control agentscurrently in use. These illustrative casehistories indicate that the risk assessmentmethodology can discriminate between agents,with some species attaining low `risk indices'and others scoring moderate or high. Riskindices should, however, not be seen asabsolute values, but as indicators to which ajudgement can be connected by biologicalcontrol experts for granting permission torelease or not.

Augmentative forms of biological control, wherenatural enemies are periodically introduced,are applied over large areas in variouscropping systems in Latin America. About 25%of the world area under augmentative control issituated in this region. Well-known examplesare the use of species of the egg parasitoidTrichogramma for management ofLepidoptera in various crops. In Mexico, forexample, about 1.5 million hectares are treatedwith Trichogramma spp. Application ofTrichogramma also occurs on large areasin Colombia and Cuba, but use is limited inother Latin American countries for economicreasons, the generally low level of educationof farmers, and, more importantly, because ofthe intensive use of pesticides that preventsuse of natural enemies. Of the other eggparasitoids, the main species used incommercial releases are Trissolcusbasalis (Wollaston) against the heteropteranNezara viridula (L.) in soybean inBrazil, and Telenomus remus Nixon againstSpodoptera frugiperda (J.E. Smith) incorn in Venezuela. Natural enemies attackinglarval and pupal stages are not used to a largeextent in augmentative biological control infield crops, with the exception of the use ofCotesia parasitoids against sugarcaneborers in Brazil and several other LatinAmerican countries. In addition to the use ofparasitoids and predators, Latin America isapplying microbial control agents on a largescale, such as viruses for control ofcaterpillars in soybean, fungi for control ofpests in coffee, cotton and sugar cane, andnematodes for control of soil pests. A recentdevelopment in biological control in LatinAmerica is the use of natural enemies andantagonists for disease and pest control inprotected cultivation, for example, inColombia, Brazil and Peru. Up to date, reliablefigures on current use of inundative andseasonal inoculative biological controlappeared hard to obtain, but it is clear thatLatin America currently is a main player in thefield of augmentative releases.

The foraging behavior of Amitus fuscipennis MacGown & Nebeker and Encarsia formosa Gahan was studied on tomato leaflets with 20 Trialeurodes vaporariorum (Westwood) larvae in the first or third stage. Ten of the whitefly larvae were previously parasitized and contained a conspecific or a heterospecific parasitoid egg or larva. The host type (host stage and/or previous parasitization) did not influence the foraging behavior of either parasitoid species. The residence time on these tomato leaflets was about 0.9 h for A. fuscipennis and 1.9 h for E. formosa. Amitus fuscipennis hardly stood still and fed little, while E. formosa showed extensive standing still and feeding. As a result, the time walking while drumming was similar for both parasitoid species. The numbers of host encounters and ovipositions per leaflet were similar for both parasitoid species. However, the residence time of A. fuscipennis was half as long as that of E. formosa so the rate of encounters and ovipositions was higher for A. fuscipennis. Amitus fuscipennis is more efficient in finding and parasitizing hosts under these conditions. The walking activity and host acceptance of the synovigenic E. formosa diminished with the number of ovipositions, but not those of the proovigenic A. fuscipennis. Encarsia formosa is egg limited, while A. fuscipennis is time limited because of its short life span and high egg load. Both parasitoid species discriminated well between unparasitized larvae and self-parasitized larvae, but discriminated poorly those larvae parasitized by a conspecific and did not discriminate larvae parasitized by a heterospecific. Self-superparasitism, conspecific superparasitism, and multiparasitism were observed for both parasitoid species. Superparasitism always resulted in the emergence of one parasitoid and multiparasitism resulted in a higher emergence of one parasitoid of the species that had parasitized first. The data suggest that A. fuscipennis is a good candidate for use in biological control of high-density spots of T. vaporariorum when we consider its high encounter and oviposition rate.

To determine the influence of pesticidetreatments on the population dynamics of thewhiteflies Trialeurodes vaporariorum(Westwood) and Bemisia tabaci Gennadius (Homoptera: Aleyrodidae)and their naturally occurring parasitoids, weperformed field experiments on insecticidesprayed and unsprayed fields during a croppingseason of snap bean in Pradera, Valle delCauca, Colombia. Substantially largerpopulations of whitefly nymphs occurred in theunsprayed field than in the sprayed field. Parasitoids were more frequent in unsprayedthan in sprayed fields with Encarsianigricephala Dozier being more prevalent thanAmitus fuscipennis MacGown & Nebeker. Insprayed fields the nymphs parasitized by E. nigricephala exceeded the unparasitizedwhitefly nymphs at the end of the croppingseason. Our results suggest that while undercurrent agricultural practices whiteflies onsnap beans cannot be exclusively controlled bynaturally occurring parasitoids, parasitoidsmay be integrated with chemical control inorder to reduce crop damage.

Amitus fuscipennis MacGown & Nebeker (Hym., Platygasteridae) is a parasitoid of the greenhouse whitefly, Trialeurodes vaporariorum (Westwood) (Hom., Aleyrodidae) on some crops as bean and snap bean ( both Phaseolus vulgaris L.) in Colombia. The searching and oviposition behaviour of A. fuscipennis was studied in the laboratory, using T. vaporariorum as a host on bean leaves. The parasitoid's basic search pathway consisted of walking, encountering the host, and drumming and probing it. While walking, the parasitoid stopped for short periods of time, partly to preen itself. Walking while searching comprised 61% of the adults' time budget and probing hosts represented 16%. After a host nymph was parasitized by A. fuscipennis, the parasitoid remained close by and continued searching for new hosts in the immediate vicinity. Such behaviour suggests area-restricted searching. The parasitoid preferred. first instars of T. vaporariorum for oviposition, resulting in long developmental times. Amitus fuscipennis had a high percentage of host acceptance resulting in a high percentage parasitism (60%) of all encountered hosts. Amitus fuscipennis, on average, walked faster before an oviposition (1.4 mm/s) than other whitefly parasitoids. The implications of these findings for the control of T. vaporariorum are discussed.

In the Mekong Delta, Vietnam, the citrus leafminer Phyllocnistis citrella (CLM) and the citrus red mite Panonychus citri are major pests in both sweet orange (Citrus sinensis) and Tieu mandarin (C. reticulata). Survey data indicate that these pest problems might be aggravated after farmers have completely destroyed the weed flora in their orchard. As citrus farmers only perceive the larger predators such as the weaver ant Oecophylla smaragdina and spiders, and have no idea about the existence of predatory mites or parasitoids, they do not know about potential positive attributes of weeds in pest management which sustain populations of natural enemies and their alternative food. IPM training programmes could use the weaver ant as an introduction to educating farmers about predatory mites and parasitoids, and should likewise emphasise the importance of beneficial asteraceous weeds such as Ageratum conyzoides. Non-crop trees such as Spondias dulcis, Mangifera indica, Eucalyptus tereticornis and Ceiba pentandra are commonly known to offer good refuge for the weaver ant. These trees should be further studied for their temporal contribution as food resource for other natural enemies of CLM and mites. Small adjustments of current weed management techniques are suggested to improve availability of pollen and nectar for beneficials at crucial moments in the cropping season, with due respect to implications at the landscape level.

A combination of biological control and host-plant resistance is needed to control greenhouse whitefly, Trialeurodes vaporariorum (Westwood). The high level of susceptibility of several host plants to whitefly, based on their performance on these plants, is well documented. These studies only provide information on the overall host-plant acceptance by whiteflies. Here, we use a method that allows an examination of the different tissue layers in the overall acceptance. The effects of plant tissue factors on whitefly probing profiles were monitored using the electrical penetration graph (EPG) method. The EPGs of whitefly originating from a culture on glasshouse cucumber, were recorded for 8 hours on sweet pepper, tomato, gerbera and cucumber plants produced in a glasshouse. On sweet pepper the graphs showed that whitefly made many short probes, had long xylem phases, short phloem phases, and the shortest duration of first probes. An opposite probing profile was found on cucumber: longer probes, shorter xylem phases, fewer phloem phases but of longer duration, and longer first probes. The values of these parameters for gerbera and tomato were intermediate. Whiteflies encountered the greatest stimulation or the least resistance in the tissues of cucumber, and the least stimulation or the greatest resistance in the tissues of sweet pepper. Rejection of host plants probably occurred before the phloem tissue was reached, as the probes prior to a whitefly leaving a host plant were so short that the stylets cannot have reached the phloem. But phloem factors also determine host-plant rejection, as phloem probing on sweet pepper - a poor host plant - was much shorter than on the other host plants. Resistance factors seem, therefore, to be located both in the epidermis/mesophyll and in the phloem. We hypothesize that the factors encountered by whitefly in the different tissue layers during probing contribute to the acceptance or rejection of a host plant. Based on the performance of whitefly on these plants, which is also reflected in the values of the EPG parameters, the order of acceptance ranked from high to low is cucumber > tomato = gerbera > sweet pepper.

Cage experiments were conducted to quantify the predation rate of the cricket Metioche vittaticollis (Sta˚l) on the eggs of rice leaffolder Marasmia patnalis Bradley. Egg predation by adult females was measured in response to changes in egg density, predator density and leaf area per cage. The number of eggs consumed per predator increased with egg density, without reaching a plateau. The predation rate decreased with increasing leaf area. The functional response could be adequately described with a linear Type I model, with the effect of leaf area included. This type of response to leaffolder egg density means that predation was not limited by prey handling time or satiation, but by the search rate. The search rate is here interpreted as the leaf area effectively searched for leaffolder eggs by a single predator in one day. Estimated search rates averaged 0.13 m2 day -1 for M. vittaticollis females. The search rate of the predators increased with prey density, but a model describing a density dependent search rate explained only 3 f the total variation in search rate. Increasing predator density per cage led to a decrease in the per capita egg predation rate when predator density was more than two per m2 leaf area. Interference might thus reduce the potential to enhance leaffolder egg predation by conservation or augmentation of predatory cricket populations

Walking speeds and walking activities (walking time divided by total time) of Trichogramma brassicae were determined at 12, 16, 20 and 25°C. Walking speed was measured during a 5-min period, and walking activity over a 4-day period. Both walking speed and walking activity were strongly influenced by temperature. Walking speed increased linearly with the temperature and was twice as high at 20 as at 12°C. At 25°C, walking activity was high during the whole day, at 20 and 16°C it decreased during the afternoon, whereas at 12°C the wasps became most active only at noon or later. At low temperatures, there was a strong individual variation in walking activity. At 25°C, T. brassicae was active most of the time, so the area searched per time unit can only increase at temperatures above 25°C if walking speed increases. At temperatures below 20°C, searching was more restricted by low walking activity than by low walking speed. Even disregarding other effects of temperature, the reduction in walking speed and walking activity at suboptimal temperatures means that T. brassicae can only search half of the area at 20°C, and only one-seventh at 15°C that it can search at 25°C.

One of the factors that may complicate biological control of the greenhouse whitefly on Gerbera jamesonii by Encarsia formosa is the rosette shape of this ornamental, which differs from the vertical shape of most vegetable plants (cucumber, egg plant, tomato, etc.). Therefore, host-habitat location and the behaviour prior to landing on uninfested and infested leaves was studied. Attraction of E. formosa from a short distance by infested leaves could not be detected: the parasitoid females landed at random on uninfested and infested leaves. After the first landing, a redistribution of the wasps occurred on the leaves. After 24 h three times as many wasps were found on the infested leaves than on uninfested ones. In a dispersal experiment with four plants, E. formosa appeared to have no preference for landing on leaves of the medium age class, which is the age class on which most of the whiteflies in a suitable stage for parasitism occur. Twenty percent of the parasitoids were found on the plants 20 min after releasing them. These results were independent of the plant cultivar and the host density on the plants. In the course of 8 h, the number of E. formosa females recovered from plants increased linearly, and this increase was greater on plants where hosts were present and also greater on the plant cultivar with the lowest trichome density. After 24 h, the percentage of females was highest (56%) on plants with the highest host density. E. formosa females were arrested on leaves where hosts were present. Contrary to our expectation, the results from the two G. jamesonii cultivars that differed strongly in leaf hairiness were not significantly different in most experiments. Only at the high host density was parasitism found to be lower on the cultivar with the higher hair density. Parasitoids may walk on top of the ‘hair coverlet’ of cultivars with high trichome density and, therefore, be hampered less than expected.

Amitus bennetti Viggiani and Evans, Encarsia formosa Gahan (two strains), Eretmocerus eremicus Rose and Zolnerowich, Eretmocerus mundus Mercet and Eretmocerus staufferi Rose and Zolnerowich, which are parasitoids of Bemisia argentifolii Bellows and Perring were compared with respect to their searching behaviour as part of a pre-introduction evaluation programme. Within a 5 cm arena, host-finding time was independent of the release distance from the host. Before oviposition, A. bennetti walked fastest, the E. formosa strains walked slowest and the Eretmocerus species intermediate. Leg length was not the most significant factor determining the differences in walking speed. After oviposition, A. bennetti and Er, eremicus had a lower walking speed and higher turn rate which is an indication of area-restricted search. The effect was strongest for A. bennetti. All species showed preference for counter-clockwise turns. Based on the walking speed alone, it is expected that A. bennetti will be the most efficient natural enemy of B. argentifolii, the Eretmocerus species intermediate and the E. formosa strains the least.

Amitus fuscipennis MacGown & Nebeker (Hymenoptera: Platygasteridae) is being evaluated as a potential biological control agent of Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) on bean crops in Colombia. The life history of this natural enemy is presented in this paper. The developmental time and longevity of fed and unfed adult parasitoids were compared under different temperature and relative humidity combinations. Mean longevity of the parasitoid in absence of hosts was highest (42.2 days) at 15°C and 75 ± 5% RH and lowest (3.9 days) at 15°C and 45 ± 5% RH. Longevity at 25°C and 75 ± 5% RH was also low (10.1 days). A. fuscipennis had the longest mean developmental time (65 days) at 15°C and 75 ± 5% RH and the shortest (23.2 days) at 25°C and 55 ± 5% RH. Combinations of moderate temperature and low relative humidity negatively affected parasitoid longevity. At constant high humidity, an increase of temperature leads to a decrease of longevity. Sex ratio was determined by (a) collecting adults in the field and rearing them to the next generation, (b) exposing different proportions of pairs of parasitoids to whitefly nymphs, and (c) examining the offspring of pairs of parasitoids. Sex ratios of populations collected in the field and reared in the laboratory were strongly female biased. Reproduction of the parasitoid is through thelytokous parthenogenesis. The species carries Wolbachia, the rickettsia that induces thelytoky. Results suggest that A. fuscipennis could be a potentially good biological control agent of T. vaporariorum in environments that are not overly dry or warm.

Summary Studies on insect reproduction in The Netherlands concern long-term projects on (a) the relationship between diapause and reproduction, (b) endocrinology and reproduction, (c) the relationship between food and reproduction, and more recently on (d) the background and evolution of parthenogenetic reproduction, (e) the link between reproduction and blood and sugar feeding in mosquitoes, and (f) the role of male sex-peptides in reproduction and mate protection. Examples of this work are presented and the importance of these fundamental studies to applied work will be discussed. At universities, curiosity-driven pure research on reproduction should have the first priority and this type of research will almost always result in applications, as we have experienced in insect pest management. Modern agriculture leads to serious environmental problems, human health problems and a dramatic reduction in biodiversity, as a result of intensive fertilizer use and frequent application of chemical pesticides. Pur...

Different types of plant resistance against the greenhouse whitefly, Trialeurodes vaporariorum, were distinguished according to the tissue location of the resistance factors. The effects of resistance factors were compared by measuring the electrically monitored and visually observed probing and feeding behaviour of whiteflies on two resistant tomato genotypes (82207 and 82216), a susceptible tomato cultivar, and a non-host plant, sweet pepper. On sweet pepper, whiteflies displayed very short first probes, very long pathway probing and spent much time on non-feeding activities such as walking and standing still. Also, a high percentage of whiteflies rejected sweet pepper without ingesting substances from the phloem vessel. These data suggest a strong resistance that is based on the factors present in surface/epidermis and/or mesophyll layers of this plant. The behaviour of whiteflies on tomato 82207 was very different from that on sweet pepper. On tomato 82207 whiteflies apparently did not perceive resistance factors on the leaf surface and in the mesophyll. Resistance factors appeared to be present in the phloem tissue, because a higher number of phloem phases, longer phloem salivation periods and shorter phloem ingestion periods were observed when compared with the susceptible tomato cultivar. Tomato 82216 showed no clear resistance factor in the phloem. The importance of combined EPG and behavioural research for host-plant resistance studies are discussed.

Populations of rice leaffolders and their natural enemies were studied in eight crops of irrigated rice in Laguna Province, the Philippines. The rice leaffolder complex consisted of three species: Cnaphalocrocis medinalis (Guenée), Marasmia patnalis Bradley and M. exigua Butler. Leaffolder population dynamics were characterized by an egg peak at maximum tillering and a broad larval peak around booting stage. Peak densities ranged from 0.2 to 2.0 larvae per hill. Most larvae originated from immigrant moths and there was no substantial second generation. The seasonal percentage egg parasitism by Trichogramma sp. ranged from 0 to 27%, and percentage larval parasitism from 14 to 56%. The braconid Macrocentrus philippinensis Ashmead was the most commonly reared larval parasitoid. Forty natural enemy taxa that may attack rice leaffolders were identified from suction and sweepnet samples: 24 predator taxa and 16 parasitoid taxa. The estimated survival rates from leaffolder egg to larval stages and between larval stages showed large variation between rice crops, but were not clearly correlated with observed levels of parasitism, natural enemy abundance, or natural enemy to leaffolder ratios. It is suggested that the generally low densities of rice leaffolders in Philippine transplanted rice are caused by their ovipositional preference for crops at the maximum tillering stage, allowing for only one generation, and by high immature mortality caused by the abundant and diverse complex of natural enemies.

Amitus bennetti Viggiani & Evans (Hymenoptera: Platygasteridae) is a recently described parasitoid of the silverleaf whitefly, Bemisia argentifolii Bellows & Perring (Homoptera: Aleyrodidae). Behaviour and life history of the parasitoid are described. The first nymphal instar of B. argentifolii is preferred by the parasitoid, but the 1st through 4th instar may be parasitised. Females first investigate hosts with their antennae, then walk over the host, and eventually step with their front legs on the leaf and insert their ovipositor inside the host facing away from the host, while the hind legs are still on the host. The time from encounter to oviposition (=latency to oviposition) is shortest on the 1st instar. Oviposition duration (meanD 39 s) comprises 50% of the handling time. Development time from egg to adult decreases from 72 days at 15C to 42 days at 20C to 28 days at 25C. We estimate that 400 degree days is required for development, with a development threshold of 10C. Adult longevity in the absence of hosts was 29, 26 and 19 days and with hosts present 8, 8 and 5 days at 15, 20 and 25C, respectively. Amitus bennetti is proovigenic and oviposits most eggs shortly after adult emergence. During the first day of their adult lives females laid 1, 31 and 49 eggs at 15, 20 and 25C, respectively. Compared with other parasitoid species, the development time of A. bennetti is very long, and the implications of this for management of B. argentifolii are discussed.

Temporal and spatial relationships between greenhouse whitefly and its parasitoid Encarsia formosa were analysed at different levels by using Rook's neighbourhood model, and for two types of greenhouses: a small greenhouse in China (15.1 m 2 ) and a large greenhouse in the Netherlands (6480 m 2 ). For whitefly eggs and larvae, there was no difference in their spatial relationship occurring within- and between-plot in the two types of greenhouses. Eggs and larvae occurrence aggregated with no special directional distributions. Whitefly adults exhibited nonlinear spatial density dependence within-plot, and moved randomly between-plot. Whitefly adults preferred to fly away in a N-S directions and to settle down at a NW-SE directions with an increase in the number of their neighbouring infected plants within-plot. The parasitoid E. formosa showed a strong relationship with the temporal and spatial distribution of the host, but influenced the spatial distribution of whitefly.

Egg mortality of rice leaf folders Cnaphalocrocis medinalisand Marasmia patnalis was studied in unsprayed irrigated rice fields in Laguna Province, the Philippines. Mortality was assessed by field exposure of laboratory-laid eggs for two days and by monitoring of field-laid eggs. Egg disappearance, the major mortality factor, was low in the first four weeks after transplanting and then increased. Egg parasitism by Trichogrammajaponicum was highest at the start of the crop and decreased to a low level towards crop maturity. Non-hatching of eggs was of minor importance. Over the total duration of the egg stage, the average disappearance of exposed laboratory-laid eggs was40%, and of field-laid eggs 46%. Egg mortality due to parasitism averaged 15% and 18%, respectively. The potential impact of egg parasitism is probably partly obscured by the disappearance of parasitized eggs. Mortality rates were highly variable between egg cohorts, but with multiple regression analysis several factors were identified that statistically explained a significant part of this variation. The results suggest that the predatory crickets Metiochevittaticollis and Anaxipha longipennis play a major role in egg disappearance, and that egg parasitism is positively dependent on the overall density of host eggs of Trichogramma in the field.

Insight into the foraging behavior of the parasitoid Encarsia formosa Gahan for whitefly hosts was gained by continuous observation of individual parasitoids on leaves of the ornamental plant Gerbera jamesonii, until females left the leaf. Comparison of the parasitoid behavior on three cultivars gave similar results. Mean searching time on uninfested G. jamesonii leaves of three cultivars was 1 h 30 min and the mean percentage of walking activity of the total observation time on those cultivars was 61%. Both parameters were not influenced by different leaf structures of Gerbera cultivars. Encounters with hosts arrested the parasitoids on the leaves. The walking activity and the percentage of host encounters that resulted in an oviposition decreased with decreasing egg load of the parasitoid. In comparison with tomato, where biological control of whiteflies is successful, only minor differences in the foraging behavior occur, except for the residence time of females, which was about three to four times longer on G. jamesonii leaves, but these leaves are about seven times larger than tomato leaves. The facts that (1) the foraging behavior of E. formosa on G. jamesonii is independant of the cultivar and (2) the foraging behavior is, in many aspects, similar to that on tomato suggest that biological control of whitefly on this ornamental plant is a potential option.

Life-history parameters of different biotypes of the whiteflyBemisia tabaci(Gennadius) species complex were reviewed. This included the B-biotype ofB. tabaci, identified asB. argentifolii(Bellows & Perring). Comparisons were made among different biotypes on cotton, among host plants for biotype B and among the whitefly speciesB. tabaciandTrialeurodes vaporariorum(Westwood), the greenhouse whitefly. The biotype identification of different populations ofB. tabaciwas summarized in a table. Biotypes discussed were A, B, Indian and biotypes of the Old World group. Temperature dependent relationships were estimated for egg development rate, development rate from egg to adult, immature mortality, adult longevity, sex-ratio, pre-oviposition period and fecundity. The fitted curves will be used as input for a simulation model of the population dynamics ofB. tabaciin a greenhouse when parasitoids are released. The model makes it possible to evaluate the integrated effect of different life-history parameters and behavioural parameters of parasitoids on whitefly population levels in a greenhouse.

Knowledge of the optimal temperatures for development and survival of biological control agents is essential for efficient mass-rearing and introduction of natural enemies in augmentative biological control programs. We studied the effect of constant and alternating temperatures on development and survival of immature stages and the sex ratio at emergence of adults of the Nearctic generalist predator Geocoris punctipes (Say). We also determined its thermal requirements. They were reared in climatic chambers at alternating (21/11°C, 24/18°C, 27/21°C and 30/26°C ± 1°C) and constant temperatures (16.8°C, 21.5°C, 24.5°C and 28.3°C ± 1°C), RH 70 ± 10% and a 14 h photophase. Survival and development of G. punctipes were the same when reared at constant and alternating temperatures. Five instars were recorded in all temperature regimes. The duration of the egg stage and each instar, as well as that of total larval development were longer, and larval survival lower when reared at 16.8°C, 21/11°C, 21.5°C and 24/18°C than at 24.5°C, 27/21°C, 28.3°C and 30/26°C. The optimal temperature range for development and survival of G. punctipes is 24.5°C to 30°C, its lower development threshold temperature is 13.5°C and its thermal constant 295.9 DD. Sex ratios were not significantly different from 1 : 1 male : female ratio in all temperature regimes. There is an excellent match between the temperature regimes at which the prey Tuta absoluta (Meyrick) and predator G. punctipes are active, which indicates that this predator will function well in crops where this pest is present.

Following a request from the European Commission, the EFSA Panel on Plant Health (PLH Panel) has been asked to deliver a scientific opinion on the risks posed by Daktulosphaira vitifoliae for the European Union (EU) and to identify and evaluate risk management options for this pest. In particular, the PLH Panel was asked to provide an opinion on the effectiveness of the current EU requirements against this organism. The PLH Panel created a working group to respond to this request by the end of May 2014. The activity will be conducted in accordance with methodologies developed by the PLH Panel and follow International Plant Protection Convention standards. Since this mandate is the first received by the PLH Panel on an important pest of Vitis sp., the first step will be to identify information gaps and additional expertise which the group might need. A major task of the working group will be to try to determine whether there are aggressive strains of D. vitifoliae outside or inside the EU territory which might pose a risk to EU grapevines.

The current methods to assess the environmental impacts of plant pests differ in their approaches and there is a lack of the standardized procedures necessary to provide accurate and consistent results, demonstrating the complexity of developing a commonly accepted scheme for this purpose. By including both the structural and functional components of the environment threatened by invasive alien species (IAS), in particular plant pests, we propose an environmental risk assessment scheme that addresses this complexity. Structural components are investigated by evaluating the impacts of the plant pest on genetic, species and landscape diversity. Functional components are evaluated by estimating how plant pests modify ecosystem services in order to determine the extent to which an IAS changes the functional traits that influence ecosystem services. A scenario study at a defined spatial and temporal resolution is then used to explore how an IAS, as an exogenous driving force, may trigger modifications in the target environment. The method presented here provides a standardized approach to generate comparable and reproducible results for environmental risk assessment as a component of Pest Risk Analysis. The method enables the assessment of overall environmental risk which integrates the impacts on different components of the environment and their probabilities of occurrence. The application of the proposed scheme is illustrated by evaluating the environmental impacts of the invasive citrus long-horn beetle, Anoplophora chinensis. (C) 2013 Elsevier B.V. All rights reserved.

The aphid predator Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is an invasive alien species in Europe and North America with negative effects on non-target species (including a decline of native ladybird populations), as well as fruit production, and human health. It is, therefore, important to find out which natural enemies could be used to reduce their numbers. Knowledge of H. axyridis’ natural enemies is summarised and data collected from the Netherlands over the past ten years are presented. Beetles were sampled from winter aggregations and from spring through to autumn with illuminated screens at night. Natural enemies were not found in samples of H. axyridis from 2003–2007. From 2008 onward H. axyridis adults were infested by: Hesperomyces virescens Thaxt. fungi (summer and winter), Parasitylenchus bifurcatus Poinar and Steenberg nematodes (winter), Coccipolipus hippodamiae (McDaniel and Morrill) mites (winter), and Dinocampus coccinellae (Schrank) parasitoids (summer and winter). Our results indicate that these natural enemies are starting to use H. axyridis as a host, but are as yet not sufficiently abundant to control the population.