生物源农药在苹果蠹蛾防治中的应用

苹果蠹蛾是世界性检疫害虫,对我国苹果优势产区构成了巨大威胁。长期依赖化学防治使该虫抗性问题变得十分严峻。为了保障食品安全,以环境友好的生物源农药替代化学农药已成为当前苹果蠹蛾防治的热点。本文对国内外现有的生物源农药,如寄生蜂、不育昆虫、颗粒体病毒、病原线虫、Bt、病原真菌、微孢子虫、性信息素、斑蝥素、多杀菌素等,在苹果蠹蛾防治中的最新应用及其存在的问题进行论述,讨论了生物源农药凭借其种类多、来源广且在用药时期上选择性强等特点,在该虫综合治理中的重要地位及面临的挑战。     

Biological tools for control of larval stages of malaria vectors – a review

The two main interventions presently being deployed for control of malaria vectors, that is, long-lasting insecticide-treated bed nets (LLINs) and indoor residual spraying (IRS) involve the use of chemical insecticides and target adult mosquitoes. Meanwhile, the potential of larval control is increasingly being acknowledged for the reduction of insecticide-resistant and/or exophagic Anopheles populations. Larval control has proven to be cost-effective and ideal for localities where mosquito-breeding sites are well defined and approachable. Utilising biological control tools to control anopheline larvae can lower the problem of resistance development, a common feature of chemical control. Fortunately, there are many options of biological larval control tools. Besides their direct impact on mortality, the effects of these tools can reach beyond the larval stage. Anopheline adults that develop from larvae exposed to biological control tools, such as entomopathogenic fungi, show reduced longevity, fecundity and susceptibility to Plasmodium infections. Combining two or more larval control tools can increase their efficacy against anopheline larvae. However, despite the identification of larval control potential in the laboratory, and in few cases in the field, many potential biological control tools have not been utilised to their fullest extent. This review provides an overview of the existing and potential biological larval control options for malaria vectors and discusses the advantages and requirements to develop them for malaria vector control.     

利用植绥螨防治烟粉虱的研究进展

对目前利用植绥螨防治烟粉虱Bemisia tabaci(Gennadius)的研究现状进行综述。主要介绍植绥螨作为生防作用物的应用历史、替代食物在利用植绥螨进行生物防治中的作用,并对代表种Typhlodromips(Amblyseius)swirskiiAthias-Henriot等作为烟粉虱生防作用物的最新研究进展、产业化现状、应用前景及可能存在的问题进行了分析,以期为这些植绥螨的引进和利用提供参考。     

Enhancing biological control of basal stem rot disease (<Emphasis Type="Italic">Ganoderma boninense</Emphasis>) in oil palm plantations

Basal Stem Rot (BSR) disease caused by Ganoderma boninense is the most destructive disease in oil palm, especially in Indonesia and Malaysia. The available control measures for BSR disease such as cultural practices and mechanical and chemical treatment have not proved satisfactory due to the fact that Ganoderma has various resting stages such as melanised mycelium, basidiospores and pseudosclerotia. Alternative control measures to overcome the Ganoderma problem are focused on the use of biological control agents and planting resistant material. Present studies conducted at Indonesian Oil Palm Research Institute (IOPRI) are focused on enhancing the use of biological control agents for Ganoderma. These activities include screening biological agents from the oil palm rhizosphere in order to evaluate their effectiveness as biological agents in glasshouse and field trials, testing their antagonistic activities in large scale experiments and eradicating potential disease inoculum with biological agents. Several promising biological agents have been isolated, mainly Trichoderma harzianum, T. viride, Gliocladium viride, Pseudomonas fluorescens, and Bacillus sp. A glasshouse and field trial for Ganoderma control indicated that treatment with T. harzianum and G. viride was superior to Bacillus sp. A large scale trial showed that the disease incidence was lower in a field treated with biological agents than in untreated fields. In a short term programme, research activities at IOPRI are currently focusing on selecting fungi that can completely degrade plant material in order to eradicate inoculum. Digging holes around the palm bole and adding empty fruit bunches have been investigated as ways to stimulate biological agents.     

昆虫群落中的集团内捕食作用

与捕食、竞争等种间关系相比 ,集团内捕食作用是一种更为复杂的种间关系 ,广泛存在于各种生态系中。该文从它的概念、类型、作用特点、与其它种间关系的区别、存在的广泛性、发生强度的影响因素、化学信息调控及其对生物防治效果的影响等几个方面介绍了天敌群落中的这一典型的种间相互关系。根据参与者在集团内捕食作用关系中的利益地位的不同 ,可将集团内捕食作用分为单向性和双向性的 ,也可分成对称性的和不对称性的。参与者的生物学特性和各种环境条件对集团内捕食作用的发生程度和结果起着重要的调控作用。在多种天敌存在的系统中 ,集团内捕食作用影响着目标生物种群动态及其生物防治的效果。深入地研究这一种间关系对增强生物控害的功能有十分重要的意义     

Cross-fertilizing weed science and plant invasion science to improve efficient management: A European challenge

Both weed science and plant invasion science deal with noxious plants. Yet, they have historically developed as two distinct research areas in Europe, with different target species, approaches and management aims, as well as with diverging institutions and researchers involved. We argue that the strengths of these two disciplines can be highly complementary in implementing management strategies and outline how synergies were created in an international, multidisciplinary project to develop efficient and sustainable management of common ragweed, Ambrosia artemisiifolia. Because this species has severe impacts on human health and is also a crop weed in large parts of Europe, common ragweed is one of the economically most important plant invaders in Europe. Our multidisciplinary approach combining expertise from weed science and plant invasion science allowed us (i) to develop a comprehensive plant demographic model to evaluate and compare management tools, such as optimal cutting regimes and biological control for different regions and habitat types, and (ii) to assess benefits and risks of biological control. It further (iii) showed ways to reconcile different stakeholder interests and management objectives (health versus crop yield), and (iv) led to an economic model to assess invader impact across actors and domains, and effectiveness of control measures. (v) It also led to design and implement management strategies in collaboration with the various stakeholder groups affected by noxious weeds, created training opportunities for early stage researchers in the sustainable management of noxious plants, and actively promoted improved decision making regarding the use of exotic biocontrol agents at the national and European level. We critically discuss our achievements and limitations, and list and discuss other potential Old World (Afro-Eurasian) target species that could benefit from applying such an integrative approach, as typical invasive alien plants are increasingly reported from crop fields and native crop weeds are invading adjacent non-crop land, thereby forming new source populations for further spread.     

利用传统生物防治控制外来杂草的入侵

随着国际贸易的日益频繁,外来有害植物入侵,严重威胁我国的自然环境和生物多样性。利用从原产地引入食性较专一的天敌来控制外来杂草是杂草生物防治的主要方式之一,有保护环境一劳永逸的效果。简要介绍了国际生物防治概况,统计表明全世界至少有133种目标杂草进行生物防治,主要分布在菊科、仙人掌科和含羞草科,63科369种无脊椎动物和真菌作为杂草生物防治的天敌,利用最多的天敌是鞘翅目象甲科和叶甲科昆虫,其中大多数项目是治理外来杂草的。杂草生物防治最活跃的国家依次为美国、澳大利亚、南非、加拿大和新西兰。重点论述了利用传统生物防治方法防治外来杂草的经典项目、国内外研究概况,以及目前面临的问题和应用前景。我国杂草生物防治起步晚,传统杂草生防的目标杂草有4种,紫茎泽兰、空心莲子草、豚草和水葫芦,其中,空心莲子草的生物防治获得成功。共引进天敌14种,输出天敌23种,与世界上生物防治先进的国家比尚有距离。中国应充分借鉴国际成功经验,对外来杂草开展生物防治。中国的生物多样性在世界上占有十分独特的地位,将在生物多样性保护中发挥重要作用。     

Optimizing nutritional conditions for the liquid culture production of effective fungal biological control agents

Spores of fungal pathogens of weeds and insects are unique in their ability to actively infect and kill their pest host. While these capabilities are advantageous in terms of their use as a contact biological control agent, or biopesticide, they also require special consideration during spore production. Directed approaches to medium optimization must consider not only spore yield but also spore qualities such as desiccation tolerance, stability as a dry preparation, and biocontrol efficacy. Nutritional conditions during culture growth and sporulation should direct the accumulation of appropriate endogenous reserves so that newly formed spores possess these advantageous qualities. Studies with the bioherbicidal fungus Colletotrichum truncatum and with the bioinsecticidal fungus Paecilomyces fumosoroseus have demonstrated the impact of nutrition on spore ‘fitness’ for use as a biological control agent. The optimization strategy used in these nutritional studies as well as a comparison of the results are presented. Received 06 February 1997/ Accepted in revised form 29 May 1997     

Conditional gene expression in invertebrate animal models

A mechanistic understanding of biology requires appreciating spatiotemporal aspects of gene expression and its functional implications.Conditional expression allows for (ir)reversible switching of genes on or off,with the potential of spatial and/or temporal control.This provides a valuable complement to the more often used constitutive gene (in)activation through mutagenesis,providing tools to answer a wider array of research questions across biological disciplines.Spatial and/or temporal control are granted primarily by(combinations of) specific promoters,temperature regimens,compound addition,or illumination.The use of such genetic tool kits is particularly widespread in invertebrate animal models because they can be applied to study biological processes in short time frames and on large scales,using organisms amenable to easy genetic manipulation.Recent years witnessed an exciting expansion and optimization of such tools,of which we provide a comprehensive overview and discussion regarding their use in invertebrates.The mechanism,applicability,benefits,and drawbacks of each of the systems,as well as further developments to be expected in the foreseeable future,are highlighted.     

我国烟粉虱主要捕食和寄生性天敌控制能力研究进展

烟粉虱Bemisia tabaci Gennadius是一种广泛分布于世界各地的多食性害虫,对我国多个省市区的农作物和观赏植物造成了严重的危害。化学杀虫剂的大量使用在杀伤天敌的同时使烟粉虱产生了不同程度的抗性,防治越来越困难;相较于化学防治,在不同地区筛选有效的天敌对烟粉虱进行生物防治是对烟粉虱进行有效控制的重要方法。我国报道的捕食性天敌共计26科109种,寄生性天敌共计2科59种。烟粉虱防治过程中由于单一的防治效果不理想,联合多个天敌资源的利用是增加对烟粉虱的生物防治效果的重要手段。本文通过对我国烟粉虱捕食性和寄生性天敌种类系统全面的整理,同时对烟粉虱优势种天敌中的单种或者多种组合利用后对烟粉虱的的控害潜能进行综述,以期为烟粉虱天敌昆虫种类的筛选和实现高效的生物防治提供科学依据。     

Quality control of Cotesia flavipes (Cameron) (Hymenoptera: Braconidae) from different Brazilian bio-factories

The goal of this study was to evaluate the quality of Cotesia flavipes from different bio-factories as biological control agents. We evaluated biological characteristics of the parasitoids throughout their lifespan, and measured the body length and width, abdomen width, thorax width and width and length of the right forewing of female and male parasitoids. Our results showed that the number of males and pupal viability were similar among the bio-factories; the number of emerged females was greater in bio-factories I and II; the egg-pupa period and the pupal period were shorter in bio-factory IV; and a greater longevity was found in bio-factories II and III. Sex ratio (at approximately 60% females) was satisfactory (in terms of suitability for release) across all bio-factories. For morphometric measurements, the body, abdomen and wing widths were similar in males; however, thorax width was greater in the males from bio-factory I; bio-factory III produced females with the highest body length; bio-factory I produced females with the greatest abdomen width; bio-factories III and IV produced females with the greatest wing length. Among the bio-factories studied, bio-factory IV produced the best quality C. flavipes, with respect to the greatest number of parasitoids per pupal mass (a mean of 57% more parasitoids) in a satisfactory sex ratio, and with the shortest developmental time, which facilitates faster rearing in the laboratory. Studies such as this, which assess the quality of a mass-produced C. flavipes, are crucial for the continued use of this parasitoid in controlling Diatraea saccharalis in the field.     

An artificial non-flying mutation to improve the efficiency of the ladybird Harmonia axyridis in biological control of aphids

The use of coccinellids in the biological control of aphids is restricted to the release of larvae because adults tend to fly away. Non-flying adults may stay longer in one place and so they and their progeny could give longer term protection to plants. This work is an attempt to produce a non-flying population by the use of a chemical mutagen and selection of adults with wing malformations through their subsequent generations. These adults are characterized by open elytra and extended wings. Some general features of this mutation were disclosed. The mutation is either unexpressed or results in malformed wings. It also seems recessive and lethal when homozygous. The adults with the mutation suffered a high level of mortality and a drastic reduction in reproductive capacity that prevents their mass rearing for biological control. This study revealed a negative relationship between wing malformations and reproductive capacity. Nevertheless, when adults with the mutation were released in greenhouses containing cucumbers infested with the aphid Aphis gossypii, they remained on the plants in higher numbers and laid eggs over a longer period of time than the control adults but their progeny were less numerous.     

Biological collections and ecological/environmental research: a review,some observations and a look to the future

Housed worldwide, mostly in museums and herbaria, is a vast collection of biological specimens developed over centuries. These biological collections, and associated taxonomic and systematic research, have received considerable long‐term public support. The work remaining in systematics has been expanding as the estimated total number of species of organisms on Earth has risen over recent decades, as have estimated numbers of undescribed species. Despite this increasing task, support for taxonomic and systematic research, and biological collections upon which such research is based, has declined over the last 30‐40 years, while other areas of biological research have grown considerably, especially those that focus on environmental issues. Reflecting increases in research that deals with ecological questions (e.g. what determines species distribution and abundance) or environmental issues (e.g. toxic pollution), the level of research attempting to use biological collections in museums or herbaria in an ecological/environmental context has risen dramatically during about the last 20 years. The perceived relevance of biological collections, and hence the support they receive, should be enhanced if this trend continues and they are used prominently regarding such environmental issues as anthropogenic loss of biodiversity and associated ecosystem function, global climate change, and decay of the epidemiological environment. It is unclear, however, how best to use biological collections in the context of such ecological/environmental issues or how best to manage collections to facilitate such use. We demonstrate considerable and increasingly realized potential for research based on biological collections to contribute to ecological/environmental understanding. However, because biological collections were not originally intended for use regarding such issues and have inherent biases and limitations, they are proving more useful in some contexts than in others. Biological collections have, for example, been particularly useful as sources of information regarding variation in attributes of individuals (e.g. morphology, chemical composition) in relation to environmental variables, and provided important information in relation to species' distributions, but less useful in the contexts of habitat associations and population sizes. Changes to policies, strategies and procedures associated with biological collections could mitigate these biases and limitations, and hence make such collections more useful in the context of ecological/environmental issues. Haphazard and opportunistic collecting could be replaced with strategies for adding to existing collections that prioritize projects that use biological collections and include, besides taxonomy and systematics, a focus on significant environmental/ecological issues. Other potential changes include increased recording of the nature and extent of collecting effort and information associated with each specimen such as nearby habitat and other individuals observed but not collected. Such changes have begun to occur within some institutions. Institutions that house biological collections should, we think, pursue a mission of ‘understanding the life of the planet to inform its stewardship’ ( Krishtalka & Humphrey, 2000 ), as such a mission would facilitate increased use of biological collections in an ecological/environmental context and hence lead to increased appreciation, encouragement and support from the public for these collections, their associated research, and the institutions that house them.     

Safety of Microorganisms Intended for Pest and Plant Disease Control: A Framework for Scientific Evaluation

Microorganisms are enormous but largely untapped natural resources for biological control of pests and diseases. There are two primary reasons for their underployment for pest or disease control: (1) the technical difficulties of using microorganisms for biological control, owing to a lack of fundamental information on them and their ecology, and (2) the costs of product development and regulatory approvals required for each strain, formulation, and use. Agriculture and forestry benefit greatly from the resident communities of microorganisms responsible for naturally occurring biological control of pest species, but additional benefits are achieved by introducing/applying them when or where needed. This can be done as (1) an inoculative release, (2) an augmentative application, or (3) an inundative application. Because of their specificity, different microbial biocontrol agents typically are needed to control different pests or the same pest in different environments. Four potential adverse effects are identified as safety issues (hazards) associated with the use of microorganisms for the biological control of plant pests and diseases. These are: (1) displacement of nontarget microorganisms, (2) allergenicity to humans and other animals, (3) toxigenicity to nontarget organisms, and (4) pathogenicity to nontarget organisms. Except for allergenicity, these are the same attributes that contribute to the efficacy of microbial biocontrol agents toward the target pest species. The probability of occurrence of a particular adverse nontarget effect of a microbial biocontrol agent may be a function of geographic origin or a specific trait genetically added or modified, but the safety issues are the still the same, including whether the microorganism intended for pest or disease control is indigenous, nonindigenous (imported and released), or genetically modified by traditional or recombinant DNA (rDNA) technology. Likewise, the probability of occurrence of a particular adverse nontarget effect may vary with method of application, e.g., whether as an aerosol, soil treatment, baits, or seed treatment, and may increase with increased scale of use, but the safety issues are still the same, including whether the microorganism is used for an inoculative release or augmentative or inundative application. Existing practices for managing microorganisms in the environment (e.g., plant pathogens,Rhizobium,plant inoculants) provide experience and options for managing the risks of microorganisms applied for pest and disease control. Moreover, experience to date indicates that any adverse nontarget effects, should they occur, are likely to be short-term or transitory effects that can, if significant, be eliminated by terminating use of the microbial biocontrol agent. In contrast, production agriculture as currently practiced, such as the use of tillage and crop rotations, has significant and long-term effects on nontarget organisms, including the intentional and unintentional displacement of microorganisms. Even the decision to leave plant pests and diseases unmanaged could have significant long-term environmental effects on nontarget organisms. Potential safety issues associated with the use of microbial biocontrol must therefore be properly identified and compared with the impact of other options for managing the pest or leaving the pest unmanaged. This paper provides a scientific framework for this process.     

Microbial selection strategies that enhance the likelihood of developing commercial biological control products

Research interest in utilizing microorganisms to create a microbial environment suppressive to plant pathogens has increased exponentially in recent years. Despite intense interest in developing biological control agents, relatively few antagonists have achieved ‘commercial product’ status. The fact that such a small proportion of active laboratory antagonists are developed into biological control products is partly due to several features common to microbial selection strategies that are widely utilized to obtain putative biological control agents: (a) relatively few candidate microorganisms are tested; (b) microbes are selected based on the results of an assay that does not replicate field conditions; and (c) the amenability of microbes to commercial development is excluded as a selection criterion. Selection strategies that enhance the likelihood of developing commercial biological control products are described. These include making appropriate choices regarding the pathosystem for biological control, the method of microbe isolation, and the method of isolate characterization and performance evaluation. A model system of developing a biological control product active against Gibberella pulicaris (Fries) Sacc. (anamorph: Fusarium sambucinum Fuckel), the primary causal agent of Fusarium dry rot of stored potatoes, is used to illustrate the proposed selection strategy concepts. The crucial importance and methodology is described, of selecting strains with enhanced potential for commercial development based on a strain exhibiting both favorable growth kinetics and bioefficacy when grown in commercially feasible liquid media. Received 06 February 1997/ Accepted in revised form 29 May 1997     

Augmentative biological control of arthropods in Latin America

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 Banker Plant Method in Biological Control

In the banker plant method, long-lasting rearing units for beneficials are created in the crop by distributing plants infested with herbivores or carrying other food items, such as pollen. The method has been widely investigated over many years and used to aid establishment, development and dispersal of beneficial organisms employed in biological control. In this review, we refine the definition of the banker plant method based on previous concepts and studies and offer the term “banker plant system” to describe the unit that is purposefully added to or established in a crop for control of pests in greenhouses or open field. The three basic elements of a banker plant system (banker plant, food source, beneficials) are discussed and illustrated with examples, and the diversity of banker plant systems (classified by target pest) used or investigated is documented. The benefits of using banker plant systems, such as low cost, increased freshness of beneficials, possibility for preventive control and for integration within IPM frameworks, make the method an interesting plant protection option with potential to enhance adoption of biological control in pest management programs.     

The application of atomic force microscopy to the study of living vertebrate cells in culture

Atomic force microscopy (AFM), a relatively new variant of scanning probe microscopy developed for the material sciences, is becoming an increasingly important tool in other disciplines. In this review I describe in nontechnical terms some of the basic aspects of using AFM to study living vertebrate cells. Although AFM has some unusual attributes such as an ability to be used with living cells, AFM also has attributes that make its use in cell biology a real challenge. This review was written to encourage researchers in the biological and biomedical sciences to consider AFM as a potential (and potent) tool for their cell biological research.     

Using parthenogenesis‐inducing Wolbachia for the selection of optimal lines of the egg parasitoid Trichogramma pretiosum for use in biocontrol

Trichogramma wasps (Hymenoptera: Trichogrammatidae) are egg parasitoids commonly employed in augmentative biological control releases against a variety of mainly lepidopteran pests. By exploiting the mechanism by which the endosymbiotic bacterium Wolbachia induces parthenogenesis in this genus, we created a set of completely homozygous Wolbachia‐infected recombinant isofemale lines (RILs), each consisting of a different combination of the genome of two well‐characterized lines of Trichogramma pretiosum Riley. We subsequently use 16 of these RILs to investigate the effect of genetic variation on various measures associated with offspring production under laboratory conditions. Unsurprisingly, substantial differences were found between the RILs in their propensity to parasitize hosts, the number of hosts they parasitize, and the levels of mortality in their offspring. Such measures can be used to choose an optimal line for biological control purposes. A method was also developed to characterize the 16 RILs using their allelic state at five loci. Essentially, this binary system uses high‐resolution melt analysis to resolve identity at each locus, with alleles originating from either the grandmaternal or grandpaternal line, and is such that each RIL can be distinguished from each other RIL by their allelic state at one or more loci. The method facilitates the easy diagnosis of line origin when two or more lines are competing with each other in competition assays, allowing for the design of more complicated tests of parasitoid quality. Future field experiments will determine which genetic line performs best under more realistic biological control conditions. The fact that these lines are infected with parthenogenesis‐inducing Wolbachia will allow for prolonged rearing without appreciable change in their genetic makeup, which should lead to a predictable biological control performance.     

The usefulness of destructive host feeding parasitoids in classical biological control: theory and observation conflict

Abstract.

• 1 We examine the conventional wisdom among biological control practitioners that destructive host feeding is a desirable attribute in parasitoids employed for classical biological control, using both the predictions of population dynamics theory and historical data on biological control introductions of Hymenoptera against Homoptera.
• 2 Population dynamics theory predicts that destructive host feeders, compared with other parasitoids, are (a) either just as likely or more likely to become established, and (b) unable to depress host equilibria as strongly.
• 3 Analyses of the B IOC AT database suggest that among parasitoids of Homoptera destructive host feeders are superior to other parasitoids with respect to both establishment rate and success rate.
• 4 We present likely explanations for the disparity between the predictions of population dynamics theory and the results of database analysis. A partial explanation for the mismatch between theory and observation with respect to the degree of pest suppression may be that females of destructive host feeding parasitoids rely less upon hosts as a food source when alternative foods such as honeydew and nectar are plentiful.
• 5 We conclude that, despite the predictions of population dynamics theory, destructive host feeders are probably better biological control agents than other parasitoids, and certainly no worse, but that it would imprudent to use destructive host feeding as the sole, or even primary, selection criterion when seeking agents for classical biological control.