Sub-lethal Effects of the Insecticides Pirimicarb and Dimethoate on the Aphid Parasitoid Diaeretiella rapae (Hymenoptera: Braconidae) When Attacking and Developing in Insecticide-Resistant Hosts

The effectiveness of parasitoids as biological control agents can be constrained by insecticide use, not only through direct mortality but also as a result of sublethal effects. Several pest aphids have become resistant to a range of insecticides and a resistant strain of Myzus persicae was used in laboratory experiments to investigate sublethal effects of the insecticides pirimicarb and dimethoate on the parasitoid Diaeretiella rapae . Both insecticides produced sublethal effects on D. rapae when the parasitoid attacked and developed in aphids that had been dipped in insecticide solutions. Dimethoate affected oviposition behaviour; females were apparently repelled by residues on the surface of dipped aphids, thus reducing their attack rate and hence the number of mummies produced. Also, the reproductive performance of parasitoids that had developed in pirimicarb-dipped aphids appeared to be adversely affected, in comparison with parasitoids that emerged from uncontaminated hosts, and this was reflected both in lower mummy production and lower attack rates. Pirimicarb, but not dimethoate, affected the sex ratio of the offspring of D. rapae that had developed in dipped aphids, causing a significant increase in the proportion of males. This only occurred when the male parent had developed in a pirimicarb-dipped aphid, suggesting that the effect involves male sterility or mating behaviour, although they appeared to mate normally. These sublethal effects are potential constraints on the efficiency and effectiveness of D. rapae as a biological control agent of aphid pests, but to assess fully their potential impact further studies need to be done using more realistic extended laboratory and semi-field techniques.     

Estimating susceptibility of biological control agents to pesticides: influence of life history strategies and population structure

In this study we examined the influences that differing life history strategies and population structures at the time of pesticide exposure have on population susceptibility to pesticides. We used life table data and a matrix projection model to incorporate combinations of mortality (lethal effect) and reductions in fecundity (sublethal effect) into estimates of intrinsic population growth rates (r) for a predator, the seven-spot lady beetle, Coccinella septempunctata L., and its prey, the pea aphid, Acyrthosiphon pisum Harris, and an aphid parasitoid, Diaeretiella rapae (M’Intosh). All three species exhibited differences in key life history variables. The aphid had the highest r and shortest generation time, the ladybeetle had the lowest r and longest generation, while the parasitoid exhibited intermediate life history characteristics. When the model was run with populations started as neonates (aphids) or eggs (lady beetle, parasitoid) for each species, ladybeetle populations were much more susceptible than either aphid or parasitoid populations 30 days after simulated exposure to a pesticide. For example, 50% mortality and a 50% reduction in fecundity resulted in a population headed toward extinction (negative r) for the ladybeetle while the parasitoid population grew exponentially (positive r) even after sustaining 70% mortality and a 70% reduction in fecundity. The aphid species maintained exponential growth after sustaining 80% mortality and an 80% reduction in fecundity. Thus, differences in life history variables accounted for the greater susceptibility of the ladybeetle to a pesticide than its aphid prey or the parasitoid over a set time interval. These differences in susceptibility were greatly reduced when the model was run starting with a mixed age/stage population (the stable age distribution) for each species indicating that population structure at the time of pesticide exposure plays a critical role in population susceptibility. These results suggest that life history attributes as well as population structure at the time of pesticide exposure both play a major role in population susceptibility to pesticides, highlighting the need to explicitly consider differences in life history variables among species when calculating compatibility of pesticides and biological control agents as well as the population structure of beneficial species at the time of pesticide application.     

Compatibility of early natural enemy introductions in commercial pepper and tomato greenhouses with repeated pesticide applications

Successful integrated pest management in protected crops implies an evaluation of the compatibility of pesticides and natural enemies (NE), as control strategies that only rely on one tactic can fail when pest populations exceed NE activity or pests become resistant to pesticides. Nowadays in Almería (Spain), growers release NE prior to transplanting or early in the crop cycle to favor their settlement before pest arrival because this improves biocontrol efficacy, although it extends pesticide exposure periods. The purpose of this research was to evaluate the compatibility of two applications of pesticides with key NE in 2‐year trials inside tomato and sweet pepper commercial greenhouses: Nesidiocoris tenuis (Reuter) (Hemiptera: Miridae), Orius laevigatus (Say) (Hemiptera: Anthocoridae) and Amblyseius swirskii (Athias‐Henriot) (Acari: Phytoseiidae). In tomato, flubendiamide and chlorantraniliprole (IOBC category 1) were compatible with N. tenuis, but chlorpyrifos‐methyl and spinosad (IOBC categories 2–3), which effectively reduced Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) density, compromised its predatory activity. In sweet pepper, chlorantraniliprole (IOBC category 1) was the only pesticide compatible with O. laevigatus while chlorantraniliprole, emamectin benzoate, spirotetramat and pymetrozine were harmless (IOBC category 1) to Amblyseius swirskii, and sulfoxaflor slightly harmful (IOBC category 2) to this phytoseiid predator.     

What makes a successful biocontrol agent? A meta-analysis of biological control agent performance

We qualitatively reviewed the biocontrol literature in two major journals, Biological Control and Environmental Entomology, over the past 10 years by scoring 878 studies into 11 biocontrol-oriented questions. Quantitative meta-analyses were then used on data from 145 studies to examine the effects of different types of biocontrol agents (parasitoids, predators, and pathogens) on several attributes of weed and pest populations. Results for our qualitative review showed that most biocontrol studies were focused on lepidopteran pests, and that parasitoids were the most common biocontrol agents used. Our quantitative review showed that, for weeds, biocontrol agents significantly reduced weed biomass (−82.0%), flower (−98.9%), and seed production (−89.4%). For pests, our quantitative review showed that biocontrol agents significantly reduced pest abundance by 130% compared to control groups, increased parasitism (+139.0%) and increased overall pest mortality (+159.0%) compared to targets not exposed to biocontrol agents. Effects on pest mortality tended to be stronger for parasitoids than predators, although reductions caused in pest abundance were much stronger when predators were used as biocontrol agents. Addition of two or more biocontrol agents increased mortality by 12.97% and decreased pest abundance by 27.17% compared to single releases. Separate sets of meta-analyses demonstrated that the negative impacts of biocontrol on non-target species were much smaller than those for target species, although adverse effects of biocontrol on non-target organisms are based on small sample sizes and should be interpreted with caution. Our results also showed that biocontrol efficacy tended to be higher when agents were generalists than when they were specialists. Large fail–safe numbers found for most of the estimated effects indicate the robustness of the results found for the efficacy of biological control programs.     

Effect of pesticides on Colpoclypeus florus (Hymenoptera: Eulophidae) and Trichogramma platneri (Hymenoptera: Trichogrammatidae), parasitoids of leafrollers in Washington.

Pesticides were evaluated for their effect on two parasitoid species, Colpoclypeus flouts and Trichogramma platneri, that are potential biological control agents of leafrollers in apple orchards. Organophosphate and carbamate insecticides were highly toxic to both parasitoids in topical applications, but foliar residues of some products were nontoxic after 7 d. At reduced rates, topically applied pyrethroids were low in toxicity to C. florus were highly toxic to T. platneri, and foliar residues were nontoxic after about 7 d. Imidacloprid and abamectin were highly toxic when applied topically to both parasitoids but were not toxic as 1-d-old residues. Insect growth regulators did not cause mortality either as topical applications or residues; however, diflubenzuron caused severe sublethal effects, completely blocking the production of C. florus offspring. Biorational pesticides, such as soap, oil, and B. thuringiensis products, caused no toxicity to C. florus but had a direct impact on T. platneri as topical applications through physical immobilization. The potential to integrate different pesticides with biological control of leafrollers and the need for a step-wise approach to evaluate the impact of pesticides against natural enemies is discussed.     

Potential for areawide integrated management of Mediterranean fruit fly (Diptera: Tephritidae) with a braconid parasitoid and a novel bait spray

The braconid wasp, Fopius arisanus (Sonan), a biological control agent for Mediterranean fruit fly, Ceratitis capitata (Wiedemann), was studied in coffee, Coffea arabica L. Fopius arisanus, comprised 79.3% of the total parasitoids (7,014) recovered from fruits collected at three small coffee farms. Data from seasonal host/parasitoid studies at a large coffee plantation also suggested that the most effective natural enemy of C. capitata in coffee may now reside in Hawaii. The original parasitoids introduced into Hawaii for C. capitata control (Diachasmimorpha tryoni (Cameron), Tetrastichus giffardianus Silvestri, and Dirhinus giffardii Silvestri) are now rare. Abundance of F. arisanus with respect to other parasitoids collected was influenced by elevation (274, 457, 610 m). Fopius arisanus was the dominant parasitoid at all three elevations, Diachasmimorpha longicaudata (Ashmead) occurred consistently, and T. giffardianus was abundant only at low elevation. The impacts on C. capitata and F. arisanus populations of bait sprays containing malathion, spinosad, or phloxine B applied to coffee were also evaluated. All three bait sprays suppressed C. capitata populations. Spinosad and phloxine B bait sprays appeared less harmful to the wasp than malathion. Fopius arisanus offers the potential for areawide management of C. capitata that includes biological control and integration with more environmentally safe chemical controls such as spinosad and phloxine B bait sprays.     

Evaluation of non-target effects of OMRI-listed insecticides for management of Drosophila suzukii Matsumura in berry crops

The spotted wing drosophila, Drosophila suzukii Matsumura, is an invasive pest of many fruit crops throughout North America, South America and Europe. The presence of this destructive pest has led to an increase in the number of insecticide applications. While conventional growers have an arsenal of different insecticides at their disposal, organic growers have a limited selection of effective options and rely heavily on applications of Entrust^®, the organic formulation of spinosad. An important part of research is to develop more tools for organic growers and evaluate the effects of insecticides intended to target D. suzukii on natural enemies in the system. The effects of six organic pesticides alone and in combination with three adjuvants and two phagostimulants were tested in laboratory bioassays on three common natural enemies in berry production systems including two predators, Chrysoperla rufilabris and Orius insidiosus, and a parasitoid wasp, Aphidius colemani. Under the IOBC toxicity rating scale, spinosad was rated consistently from slightly harmful to harmful across natural enemy species and residue age (the effects of pesticides over time). Sabadilla alkaloids caused mortality to O. insidiosus equal to that of spinosad. All tested pesticides were at least slightly harmful to A. colemani, and the adjuvant polyether-polymethylsiloxane-copolymer polyether caused mortality that was not significantly different from spinosad. In general, neither the addition of adjuvants nor phagostimulants increased the mortality of the insecticides tested. The exception was polyether-polymethylsiloxane-copolymer polyether, but it is unclear whether it increased the toxicity of the pesticides or was simply toxic itself since it caused high mortality to A. colemani when applied alone. Sublethal effects were measured for two predatory species by measuring eggs laid and % egg hatch. Minimal sublethal effects were observed in C. rufilabris. In contrast, all tested insecticides caused reduced egg hatch in O. insidiosus compared with the control.     

Impacts of biological control and invasive species on a non-target native Hawaiian insect

The potential for classical biological control to cause unintended harm to native species was evaluated in the case of the endemic Hawaiian koa bug, Coleotichus blackburniae White (Hemiptera: Scutelleridae), and parasitoids introduced to Hawaii for control of an agricultural pest, the southern green stink bug, Nezara viridula (L.) (Hemiptera: Pentatomidae). Parasitism of C. blackburniae eggs, nymphs and adults by biocontrol agents was quantified across a wide range of habitats and compared to other sources of mortality. Egg mortality due to the biocontrol agent Trissolcus basalis Wollaston (Hymenoptera: Scelionidae) was low (maximum 26%) and confined to elevations below 500 m on a single host plant. Predation, mainly by alien spiders and ants, was the greatest source of egg mortality (maximum 87%). Parasitism of adult C. blackburniae by the biocontrol agent Trichopoda pilipes (F.) (Diptera: Tachinidae) was near zero at 21 of 24 sites surveyed. Three sites with high bug density had higher levels of T. pilipes parasitism, reaching maxima of 70% among adult female bugs, 100% among males and 50% among fifth instars. Male-biased parasitism indicated that T. pilipes is adapted to using male aggregation pheromone for finding C. blackburniae hosts. The relative impacts of biocontrol agents and other sources of mortality were compared using life tables. Invasive species, particularly generalist egg predators, had the greatest impacts on C. blackburniae populations. Effects of intentionally introduced parasitoids were relatively minor, although the tachinid T. pilipes showed potential for large impacts at individual sites. In retrospect, non-target attacks by biological control agents on C. blackburniae were predictable, but the environmental range and magnitude of impacts would have been difficult to foresee.     

Male mating preference of two cryptic species of the herbivorous insect Eccritotarsus catarinensis

When using biological control against pest populations, more than one biocontrol agent might be introduced simultaneously. This could be counterproductive in the event of negative interactions between the biocontrol agents. Within and between species interactions have a strong impact on mating behaviour and reproduction, and can have an impact on the effectiveness of biological control. We studied the reproductive compatibility between two geographically isolated strains (Brazil and Peru) of Eccritotarsus catarinensis (Heteroptera: Miridae), a biocontrol agent against the invasive aquatic weed, Eichhornia crassipes. By performing inter- and intra-species mating experiments, we investigated whether or not males from each of the cryptic species would be able to distinguish between partners from either species, and if they would mate with partners from the opposite species. Our results showed the decrease in lifetime fecundity, and most importantly, the lack of production of offspring from eggs resulting from forced hybridisation. We showed that Peruvian males mated only with females from their own species, and did not mate with females from the Brazilian species. In contrast, Brazilian males mated equally with females from both species, but needed significantly more time in order to commence a mating, and no offspring were produced from eggs resulting from hybridisation. Although future studies demand more rigorous controls, our results indicate asymmetrical sexual isolation between the two species. We speculate on mechanisms involved in reproductive isolation in the two cryptic species, and the possible implications for effective biocontrol, and include some morphological measurements that might support our assumptions.     

Sublethal effects of fenpyroximate on life table parameters of the predatory mite <Emphasis Type="Italic">Phytoseius plumifer</Emphasis>

Knowledge of the lethal and sublethal effects of pesticides on biological control agents is necessary for successful implementation of integrated pest management programs. The sublethal effect of acaricide fenpyroximate (Ortus^®) on performance of females and their offspring of Phytoseius plumifer (Canestrini &; Fanzago), an efficient predator of Tetranychus urticae Koch, was determined under laboratory conditions. The adult predators were exposed to the residues of the acaricide on fig leaves and the LC₅₀ value was determined based on a concentration–response analysis. The results showed that sublethal concentrations of fenpyroximate significantly affected the fecundity and longevity of the treated females of P. plumifer. Reproductive and life table parameters of the subsequent generation were also affected by sublethal concentrations. It could be concluded that sublethal concentrations can significantly reduce the population growth of P. plumifer and this should be considered in integrated pest management programs.     

After biocontrol: Assessing indirect effects of insect releases

Development of biological control agents for weeds has been motivated by the need to reduce the abundance and distribution of a pest plant where chemical and mechanical control were not cost effective. Primary objectives have been direct reduction in abundance of the target and, secondarily, the increase of desirable species. Recently, wildland weeds have become a focus of biological control projects. Here, desired outcomes include both reduction of the target and indirect effects of increased diversity and abundance of native species and restoration of ecosystem services. However, goals and benefits of biocontrol programs are not always well-articulated and direct and indirect impacts are not easily predicted. We evaluated the extent to which several successful biological control projects for weeds of rangelands and waterways measured indirect impacts on invaded ecosystems. We also examined biocontrol of a wildland pest tree for which the principal objective is restoration of ecosystem services. We found few quantitative assessments of the impacts of pest plant reduction on community composition or ecosystem processes. All examples documented variation in the impacts of agent(s) across the invasive range of the target plant as well as variation in impacts on the invaded ecosystem. However, without appropriate quantitative information, we cannot evaluate site characteristics that may influence vegetation responses. Most successful weed management programs integrated the use of biocontrol agents with other weed management strategies, especially modifications of disturbance and competing vegetation. Discussion and evaluation of responses of nontarget species would improve our understanding of the context-specificity of outcomes.     

Why demographic and modeling approaches should be adopted for estimating the effects of pesticides on biocontrol agents

Recent studies have shown that simplistic measures of toxicity such as the LC₅₀ do not provide enough information about the actual effects that may occur in pesticide-exposed populations over longer time periods than a few days. In this paper we discuss the use of demography and population modeling for estimation of pesticide effects on pest and beneficial species and argue that these new approaches are essential to further our understanding of the potential impacts that pesticides might have on both pest and beneficial species such as biological control agents.     

Effects of neem-treated aphids as food/hosts on their predators and parasitoids

Abstract:  Experiments were conducted in the laboratory and in greenhouses. Of three neem preparations sprayed upon eggs, only neem oil (NO) exerted a negative impact on the hatching rate of Coccinella septempunctata and Chrysoperla carnea . First instar larvae of Episyrphus balteatus proved to be highly susceptible, when feeding 24 h on aphids sprayed with neem kernel water extract (NKWE). First instar larvae of C. septempunctata showed a very high mortality when feeding on aphids sprayed with different neem preparations. Aphid feeding and live span was reduced. When NKWE had been applied to the soil, the mortality of larvae of E. balteatus and C. septempunctata were lower, when feeding on aphids. Second instar larvae of C. septempunctata were far less susceptible when feeding 48 h on neem-sprayed aphids than first instars; the time of their development was prolonged, and aphid consumption reduced. Larvae of C. carnea proved to be less susceptible, when feeding on neem-sprayed aphids, than E. balteatus and C. septempunctata . In C. carnea , however, significant influences were also observed in aphid consumption, time of development, mortality, longevity, and rate of deformity. NO, containing a very low concentration of azadirachtin A, had stronger negative effects than NeemAzal-T/S^®, in all observations. In the parasitoid Diaeretiella rapae , NKWE application to the soil induced negative reactions, when aphids on these plants were parasitized: low percentage parasitization, lowered mummy weight, low emergence rate of adults of F1 and even of F2. Foliar sprays of NKWE had less severe effects in this parasitoid species. The results are discussed with regard to their theoretical and practical significance.     

Evaluation of Fopius arisanus as a biological control agent for the olive fruit fly in California

1 The egg‐prepupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) was evaluated in quarantine facilities as a potential biological control agent for the olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California, U.S.A. 2 Nontarget testing of two weed biological control agents confirmed that F. arisanus will not attack Tephritidae that feed in inflorescences or galls. It may, however, pose risks to native Tephritidae that feed in fruit. 3 Females preferentially oviposited in eggs, although first‐instar B. oleae were also attacked. Low lifetime reproductive potential and high rates of direct mortality inflicted on host eggs indicate that rearing on B. oleae may prove difficult. 4 In multiparasitized B. oleae, F. arisanus prevailed in competition against two species of larval–pupal parasitoids, Diachasmimorpha kraussii and Psyttalia concolor (both Hymenoptera: Braconidae). 5 The broad host‐range of F. arisanus with respect to fruit‐feeding Tephritidae may preclude its introduction to California, as may its low fecundity and its intrinsic competitive superiority over larva l–pupal parasitoids, which include specialists on B. oleae that are currently being introduced to California. High rates of direct mortality, however, point to potential uses in augmentative biological control. Whether or not F. arisanus is released in California, its biology as a parasitoid of B. oleae has been little studied to date and the results herein may be applied in other regions worldwide where B. oleae is a problem.     

Juvenile and sublethal effects of selected pesticides on the leafminer parasitoids Hemiptarsenus varicornis and Diglyphus isaea (Hymenoptera: Eulophidae) from Australia

The pest leafminers Liriomyza huidobrensis (Blanchard), Liriomyza sativae (Blanchard), and Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) have spread into South East Asia and Oceania, and they are likely to reach Australia in the near future. Two translaminar pesticides, cyromazine and abamectin, currently provide effective chemical control of these pests, but because parasitoids can play an important role in controlling and preventing leafminer outbreaks, understanding the impact of pesticides on leafminer parasitoids is vital. Here, we tested larval and pupal mortality and sublethal effects of abamectin, cyromazine, and the widely used fungicide mancozeb on two common Australian leafminer parasitoids, Hemiptarsenus varicornis (Girault) and Diglyphus isaea (Walker). Abamectin caused significant mortality to larvae and pupae of both parasitoid species but cyromazine and mancozeb did not. Progeny production and longevity of H. varicornis were not affected by adult exposure to cyromazine and mancozeb, nor did direct pupal exposure decrease number of progeny produced by either parasitoid. Mortality of H. varicornis females emerging from leaves treated with abamectin was high for up to 72 h after eclosion but those surviving beyond 72 h did not differ from control females in the number of progeny produced. Mancozeb did not influence leaf residence time or parasitism by H. varicornis females. Cyromazine and the fungicide mancozeb were concluded to be compatible with the parasitoids tested and suitable for integrated pest management of leafminers should outbreaks of pest species occur in Australia. Abamectin should be used with caution because it caused significant mortality in both parasitoids tested here.     

Disruptive Sublethal Effects of Insecticides on Biological Control: Altered Foraging Ability and Life Span of a Parasitoid after Feeding on Extrafloral Nectar of Cotton Treated with Systemic Insecticides

Predictions in integrated pest management on the compatibility of an insecticide with biological control often are based on incomplete screening tests. While measuring levels of mortality from direct insecticide exposure is a very common screening method, possible sublethal effects as a result of either direct or indirect insecticide exposure remain relatively unknown. The impact of sublethal effects on the success of biological control can be as deleterious as mortality. Here, we report the reduced host foraging ability and longevity of the parasitoid Microplitis croceipes Cresson (Hymenoptera: Braconidae) after feeding on extrafloral nectar from cotton (Gossypium hirsutum L., Malvaceae) plants that were treated with systemic insecticides. The insecticides used in this study are regularly applied in cotton-growing areas in the United States. For all tested insecticides, longevity of M. croceipes females that fed on nectar from cotton was affected for at least 10 days after plants were treated with insecticides. Moreover, the parasitoid's host foraging ability was severely affected for periods ranging from 2 days (imidacloprid) to 18 days (aldicarb) after insecticide application. The consequences of these sublethal effects on the success of biological control are discussed.     

Genetic analysis of an introduced biological control agent reveals temporal and geographic change,with little evidence of a host mediated shift

The evolution of introduced biological control agents is largely un-explored. Although much is theorized, there is little empirical evidence quantifying the evolutionary dynamics of a biocontrol agent after release into a new environment. In this study we use Diachasmimorpha tryoni, a purposefully introduced biocontrol agent of Ceratitis capitata, to model and quantify spatial, temporal, and host-related evolutionary patterns. This parasitoid has undergone a host shift in its introduced environment, Hawaii, to the gall forming weed biocontrol agent, Eutreta xanthochaeta, an interaction likely mediated by competition for C. capitata with the egg-larval parasitoid Fopius arisanus. To elucidate potential evolutionary patterns we analyzed microsatellites and sequence data extracted from Hawaii and Australian population clusters defined by Structure, in Genepop, Canoco, and IBDWS. Our analysis revealed structuring of Hawaiian D. tryoni populations as defined by significant historic influences related to temporal structure, geographic space, host guild, and augmentative releases. The host-shift parasitoids were not genetically distinct from other Hawaii populations. There were small changes in microsatellite DNA at the population level, but only between Australia and Hawaii populations, not at the host level. These results show that D. tryoni has not undergone host-mediated evolution since introduction to Hawaii, despite the fact that they have expanded their host range in Hawaii to include the gall-forming E. xanthochaeta. To our knowledge this is the first study to quantify genetic differentiation of a biological control agent over geographic space and time using contemporary and museum specimens.     

Effects of floral resources on the efficacy of a primary parasitoid and a facultative hyperparasitoid

Resources added to agroecosystems to enhance biological control are potentially available to multiple members of the resident insect community—not only the biological control agents for which the resources are intended. Many studies have examined the effects of sugar feeding on the efficacy of biological control agents. However, such information is lacking for other, interacting species such as facultative hyperparasitoids, which may contribute to pest suppression but can also interfere with introduced biological control agents. Under greenhouse conditions, we tested the direct effects of sugar and nectar provisioning on the longevity, host‐killing impact and offspring production of two pupal parasitoids associated with leek moth, Acrolepiopsis assectella: the introduced biological control agent, Diadromus pulchellus, and the native facultative hyperparasitoid, Conura albifrons. Adding sucrose, buckwheat or a combination of buckwheat and common vetch to a sugar‐deprived system (potted leek plants in cages) increased parasitoid longevity and resulted in higher leek moth parasitism and mortality compared to water or common vetch treatments. However, the two parasitoid species exhibited a distinct temporal response to the treatments, likely influenced by differences in their life histories. This study provides insight into how integrating conservation biological control techniques could affect the success of a classical biological control programme.     

Development of a biological control-based Integrated Pest Management method for Bemisia tabaci for protected sweet pepper crops

The tobacco whitefly, Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae), is a key pest in commercial sweet pepper crops in southeast Spain. Its biological control is currently based on augmentative introductions of the parasitic wasp Eretmocerus mundus Mercet (Hymenoptera: Aphelinidae), which need to be occasionally supplemented with pesticide applications. These pesticides can be harmful for the biological control agents. Therefore, it is important to improve the current strategy by reducing dependency on pesticides. Two potential solutions are conceivable: addition of another effective biocontrol agent or application of pesticide prior to the release of biocontrol agents. The mirid bug Nesidiocoris tenuis Reuter (Heteroptera: Miridae) and the predatory mite Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae) are promising candidates as additional biocontrol agents. The aim of the present study was to test these possible solutions in two subsequent trials, i.e., a 'selection' and an 'improvement' experiment. In the selection experiment, four treatments were compared: E. mundus , N. tenuis  +  E. mundus , A. swirskii  +  E. mundus , and A. swirskii  +  N. tenuis  +  E. mundus . Amblyseius swirskii appeared able to significantly increase effectiveness against the pest, in contrast to N. tenuis, which did not contribute to whitefly control. The best strategy was the combination of E. mundus and A. swirskii . In the improvement experiment, three treatments were compared: E. mundus , A. swirskii  +  E. mundus , and A. swirskii  +  E. mundus +  pesticides. Amblyseius swirskii again proved capable of significantly reducing whitefly populations, and the implementation of pesticides before the release of the biocontrol agents was shown to increase the effectiveness against the pest even more.     

Native insects as targets and non-target species of biological control

The use of population models for predicting desirable and undesirable outcomes of biological control are described using three case studies from New Zealand. The first reviews the models of Barlow and colleagues used to understand and improve the control of native grass grub Costelytra zealandica populations by augmentative application of pathogenic Serratia entomophila bacteria. A variety of modelling approaches have been used to predict grass grub outbreaks and thus the cost-effectiveness of applying bacteria. Models have also been developed to understand the competitive interactions between pathogenic and non-pathogenic forms of the bacteria. The other two case studies show how retrospective modelling has been used to quantify the non-target impact of introduced biological control agents. The parasitoid Microctonus aethiopoides was introduced to control the lucerne pest Sitona discoideus, but was found to disperse outside of the target habitat and attack several native weevil species in the Entiminae family. Retrospective modelling suggests that, given average parasitism levels of 15%, native Irenimus spp. and Nicaeana spp. weevil populations are likely to have been reduced by 8% due to non-target parasitism. Similarly, population models have shown that native red admiral butterfly (Bassaris gonerilla) populations are likely to have been reduced by 5% due to non- target parasitism by Pteromalus puparum, which was introduced to New Zealand for the control of the cabbage white butterfly (Pieris rapae).