The ecological barriers to the recovery of bridal creeper (Asparagus asparagoides (L.) Druce) infested sites: Impacts on vegetation and the potential increase in other exotic species

Abstract To protect native biodiversity from environmental weeds, the impacts that these weeds cause need to be known before weed control commences. Asparagus asparagoides (L.) Druce (bridal creeper) (Asparagaceae) is a serious environmental weed and has been selected for biological control in Australia. To predict the responses of plant communities to the control of bridal creeper, a prerelease baseline of the impacts of bridal creeper on native plant communities was undertaken. Plant assemblages in areas invaded by bridal creeper were compared with reference areas that contained little or no bridal creeper. Areas invaded by bridal creeper contained 52% fewer native plant species when compared with nearby reference areas. However, there was no difference in the number of other exotic plant species between areas. Similar trends were found for the germinable seed bank. Although a greater number of exotic species were present in the seed bank compared with the vegetation surveys, there was still no difference between areas with and without bridal creeper. In a glasshouse trial, exotic species germinated more frequently compared with native species. This could indicate that as bridal creeper density decreases following control, exotic species have an advantage over native species when colonizing areas left vacant by bridal creeper. Second, as bridal creeper areas contained reduced native species richness and cover, they may be susceptible to further weed invasion after bridal creeper is removed. Therefore, simply reducing the presence of bridal creeper may not guarantee successful restoration of invaded areas and additional restoration efforts will be needed to ensure the ultimate goal of protecting native biodiversity is reached.     

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.     

Responses of invertebrates to herbicide in Salix cinerea invaded wetlands: Restoration implications

The use of herbicides to control weeds, particularly large invasions, has now become an essential management tool in many ecological restoration projects. The herbicide glyphosate is routinely used to control the invasive weed, Grey Willow (Salix cinerea), within New Zealand wetlands. However, little is known about the effects of glyphosate on invertebrates. We determine the short‐term effects of glyphosate on the abundance and composition of the nontarget canopy invertebrate community in wetlands invaded by Grey Willow in New Zealand. Initially, the application of glyphosate and a surfactant showed no detectable effect on the canopy invertebrates examined in this study. However, 27 days after herbicide application, significant Grey Willow canopy loss caused dramatic decreases in the abundance of invertebrates in the glyphosate‐treated plots compared with the unsprayed plots. Invertebrates appeared to be sensitive to changes in vegetation structure, such as canopy loss. These results agree with previous studies that have shown that the negative impacts of glyphosate on invertebrate communities are related to indirect effects via habitat modification as the herbicide‐treated vegetation dies. From a terrestrial invertebrate perspective, this study suggests that the use of glyphosate herbicide is suitable for the control of invasive weeds within wetland restoration projects as it appears to have negligible impact on the canopy invertebrate assemblage.     

Apparent competition can compromise the safety of highly specific biocontrol agents

Despite current concern about the safety of biological control of weeds, assessing the indirect impacts of introduced agents is not common practice. Using 17 replicate food webs, we demonstrate that the use of a highly host-plant specific weed biocontrol agent, recently introduced into Australia, is associated with declines of local insect communities. The agent shares natural enemies (predators and parasitoids) with seed herbivore species from native plants, so apparent competition is the most likely cause for these losses. Both species richness and abundance in insect communities (seed herbivores and their parasitoids) were negatively correlated with the abundance of the biocontrol agent. Local losses of up to 11 species (dipteran seed herbivores and parasitoids) took place as the biocontrol agent abundance increased. Ineffective biocontrol agents that remain highly abundant in the community are most likely to have persistent, indirect negative effects. Our findings suggest that more investment is required in pre-release studies on the effectiveness of biocontrol agents, as well as in post-release studies assessing indirect impacts, to avoid or minimize the release of potentially damaging species.     

Integrating Management Techniques to Restore Sites Invaded by Mile‐a‐Minute Weed,Persicaria perfoliata

Efforts to suppress an invasive weed are often undertaken with the goal of facilitating the recovery of a diverse native plant community. In some cases, however, reduction in the abundance of the target weed results in an increase in other exotic weeds. Mile‐a‐minute weed (Persicaria perfoliata (L.) H. Gross (Polygonaceae)) is an annual vine from Asia that has invaded the eastern United States, where it can form dense monocultures. The host‐specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the United States in 2004 as part of a classical biological control program. At three sites invaded by mile‐a‐minute weed, biological control was integrated with pre‐emergent herbicide use and two densities of native plantings. After 2 years, native plant cover differed significantly and was greater than 80% in the plots with plantings and pre‐emergent herbicide but less than 30% in the planting treatments without herbicide. Where mile‐a‐minute cover decreased at the two sites with the greatest pressure from exotic plants, plots were dominated by another exotic weed, Microstegium vimineum (Trin.) A. Camus, Japanese stiltgrass. The combination of biocontrol, pre‐emergent herbicide, and revegetation with native plants suppressed mile‐a‐minute weed, prevented invasion by Japanese stiltgrass, and increased the abundance of native plants. The selection of the management strategies used to control mile‐a‐minute weed determined the extent of recovery of the native plant community.     

Success and safety in the biological control of environmental weeds in New Zealand

Biological control, using specialist insect herbivores and plant pathogens, can be a self‐sustaining, cost‐effective and low‐risk tool for the management of environmental weeds. Agents have been recorded attacking non‐target plants in New Zealand and elsewhere, but the effects are usually minor and/or transitory. It seems probable that only two cases, worldwide, will result in significant damage to non‐target plants (representing 0.5% of the nearly 400 insect, mite, or fungal species used in classical weed biocontrol). Both of these cases were predictable from host range testing. Negative indirect, or ‘downstream’, ecological effects from specific weed biocontrol agents are difficult to predict and measure. They are probably insignificant compared to the impacts of the invasive plants that the agents are introduced to control. However, it is necessary to balance the risks associated with any introduction against the environmental benefits from controlling a weed to a predicted level. Recent analyses suggest that success rates are better than generally perceived. For New Zealand programmes, where enough time has lapsed to allow assessment, we calculate a full/partial success rate of 83%. Many of the costs associated with environmental weeds are difficult to quantify. Detailed risk assessment will make biological control programmes more expensive and time‐consuming, so that reliance on non‐biological management methods for environmental weeds may actually increase. The costs of biocontrol programmes against some New Zealand weeds can be kept down by using research already carried out in Australia and other countries, and the process is reciprocal. Developing international consortia of sponsors is also a potential way to fund programmes against weeds shared by several countries.     

Limited Effects of Large‐Scale Riparian Restoration on Seed Banks in Agriculture

I examined the effect of riparian forest restoration on plant abundance and diversity, including weed species, on agricultural lands along the Sacramento River in California (United States). Riparian forest restoration on the Sacramento River is occurring on a large‐scale, with a goal of restoring approximately 80,000 ha over 160 km of the river. In multiuse habitats, such as the Sacramento River, effects of adjoining habitat types and movement of species across these habitats can have important management implications in terms of landscape‐scale patterns of species distributions. Increased numbers of pest animals and weeds on agricultural lands associated with restored habitats could have negative economic impacts, and in turn affect support for restoration of natural areas. In order to determine the distribution and abundance of weeds associated with large‐scale restoration, I collected seed bank soil samples on orchards between 0 and 5.6 km from adjacent restored riparian, remnant riparian, and agricultural habitats. I determined the abundance, species richness, and dispersal mode of plant species in the seed bank and analyzed these variables in terms of adjacent habitat type and age of restored habitat. I found that agricultural weed species had higher densities at the edge of restored riparian habitat and that native plants had higher densities adjacent to remnant riparian habitat. Weed seed abundance increased significantly on walnut farms adjacent to restored habitat with time since restored. I supply strong empirical evidence that large areas of natural and restored habitats do not lead to a greater penetration of weed species into agricultural areas, but rather that weed penetration is both temporally and spatially limited.     

Exotic wildland weeds serve as reservoirs for a newly introduced cole crop pest,Bagrada hilaris (Hemiptera: Pentatomidae)

The role of wildland weeds that serve as alternate hosts for insect pests has not been directly examined for the potential to sustain pest populations or facilitate pest outbreaks. The development of weed biological control programmes is also complicated by weed species that are closely related to economically important or native plants, especially rare or special status species. In recent surveys in southern California, USA, we found a newly introduced insect pest of cole crops, Bagrada hilaris Burmeister (Bagrada bug), feeding on Brassicaceae weeds in riparian areas adjacent to agricultural fields where cole crops are routinely grown. Insect populations grew to levels well over action thresholds and caused severe damage to populations of the invasive weed, Lepidium latifolium (perennial pepperweed). The numerical response of B. hilaris on L. latifolium and other Brassicaceae weeds in natural areas may pose a significant challenge to effectively managing pest populations in crops. However, the accidental introduction of this insect provides the opportunity to examine plant–insect interactions with important implications for development of biologically based control methods for weeds.     

Grasses as appropriate targets in weed biocontrol: is the common reed, <Emphasis Type="Italic">Phragmites australis</Emphasis>, an anomaly?

Despite their importance as invasive species, there has been a hesitation to target grasses in classical biocontrol. This historic bias appears to be changing with multiple active research and release programs. Similarly, biocontrol workers appear to avoid targeting species with native congeners. These biases appear inappropriate as the ecological and entomological literature provide abundant evidence for sub-genus specificity for many herbivores, including those attacking grasses. The biocontrol program targeting Phragmites australis (Cav.) Trin. ex Steud (Poaceae) provides an informative example with endemic subspecies in North America and many sub-genus specific herbivores, including potential European control agents. Grasses and target weeds with congeneric native species require rigorous host range testing, similar to all other targets in current weed biological control programs. Furthermore, it appears prudent to ask petition reviewers and regulatory agencies to abandon their focus on results of no-choice studies and to distinguish between trivial feeding and demographic impacts.     

Biological control of weeds: an analysis of introductions,rates of establishment and estimates of success,worldwide

The foremost document that comprehensively reports on biological control introductions against weeds—‘Biological control of weeds: a world catalogue of agents and their target weeds’—has been updated and now includes all deliberate releases made through 2012. It includes data on 1555 intentional releases of 468 biological control agent species used against 175 species of target weeds in 48 plant families, in 90 countries. For 55 (31.4%) of the target weed species, only one biocontrol agent was introduced. The largest number of agent species (44) was introduced for the biological control of Lantana camara (Verbenaceae). Three insect orders (Coleoptera, Lepidoptera and Diptera) comprised about 80% of all biocontrol agent species released and releases made. Of the 468 biocontrol agent species introduced, 332 (70.9%) established in at least one instance. Of the 313 species, for which impact could be categorized, 172 (55.0%) caused medium, variable or heavy levels of damage (impacts). Of all releases made through 2012, 982 (63.2%) led to establishment. Forty-two releases were judged too early post-release to categorize impact, leaving 940 releases for which impact analyses were conducted. Similar to agent species, approximately half of the established releases (503 or 53.5%) caused medium, variable or heavy levels of damage on the target weeds, and almost a quarter of releases (225 or 23.9%) caused heavy impact. Across all countries and regions, 65.7% of the weeds targeted for biological control experienced some level of control. These data indicate the value of this practice, on its own, or as a supplement to other methods, in the management of invasive plants.     

A proposal to use plant demographic data to assess potential weed biological control agents impacts on non-target plant populations

Weed biocontrol programs aim to reduce the spread and population growth rate of the target plant while stabilizing or increasing populations of those native species considered under threat by invasive plants. This goal is not unique to weed biocontrol but applies to all other invasive plant management techniques, though such information is rarely collected. Without this information, success of management interventions can be ambiguous, and regulatory agencies, the public, policy makers, funders and land managers cannot be held accountable for chosen treatments. A fundamental reform, including use of demographic studies and long-term assessments, are essential to guide weed biocontrol programs. We propose to add use of plant demography (an assessment of how environmental factors and ecological interactions, for example competition, disease or herbivory, may affect plant populations by altering survival, growth, development and reproductive rates of plant individuals) during host specificity risk assessments of potential biological control agents. Demographic models can refine assessments of potential impacts for those plant species that experience some feeding or larval development during host specificity testing. Our proposed approach to focus on impact on plant demography instead of attack on plant individuals is useful in appropriately gauging threats potential weed biocontrol agents may pose to non-target species after field release.     

Prerelease efficacy assessment, in quarantine, of a tephritid gall fly being considered as a biological control agent for Cape-ivy (Delairea odorata)

Due to the long-standing emphasis on only releasing host-specific agents, classical biological control of weeds has an enviable track record of few direct impacts to nontargets. However, even an agent whose host-range is restricted solely to the target weed can have indirect impacts. Such indirect impacts are most likely if, after release, the populations of the agent build up to high numbers without causing accompanying declines in the populations of the target weed. Therefore, it is advisable, prior to release, to demonstrate that the candidate agent is not only host-specific, but that it has clear potential to depress populations of the target weed. Prerelease efficacy assessments (PREA) of potential weed biocontrol agents are not yet common, and are most easily done in the region where both the target and the potential agent are native. We present an example of a PREA performed under strict containment conditions of an approved quarantine facility. A gall-forming fly, Parafreutreta regalis, from South Africa is being considered for release in California to control Cape-ivy, Delairea odorata. We conducted two trials exposing test Cape-ivy plants to two different densities of this fly, and, after approximately two months, comparing the growth of the galled vines to similar vines that had not been exposed to flies. Under both the high density (10 pairs of flies/plant) and low density (2 pairs/plant) treatments, the galled vines exhibited visible stunting, and the ungalled stems were longer, and had more nodes and larger leaves. These trials confirmed that relatively subtle, sublethal impacts on the target can be quantified, even under strict containment conditions, and this should encourage others to assess, prior to release, the potential impact of prospective agents on their proposed target.     

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。     

Evaluating the post-release efficacy of invasive plant biocontrol by insects: a comprehensive approach

We propose a comprehensive program to evaluate the post-release phase of biocontrol programs that use insect herbivores to control invasive plant species. We argue that any release should be done in randomized release and non-release sites and should be followed up by well-replicated sampling and experimental protocols that evaluate the degree of success or failure. These follow-up studies should include landscape scale monitoring across relevant habitat gradients of (1) the abundance of the biocontrol agent, (2) the impact of the biocontrol agent on the target plant species, (3) the potential for non-target effects, and (4) the response of native species and communities to a reduction in the invasive species. We also argue that (5) experimental reductions of the biocontrol agent are required to eliminate the chance that the putative impact of the biocontrol agent is not confounded with other causes. Finally, we describe six scenarios, informed largely by a community ecology perspective, in which a biocontrol agent may decrease the abundance or vigor of the target plant species but not lead to successful control where native communities re-establish. We classify these failure scenarios as either direct or indirect effects of the invasive plant species: Native Source Limitation, Static Competitive Hierarchies, Novel Weapons, Trophic Shifts, Invasive Engineering and Associated Invasives. Overall, we argue that well replicated and landscape-scale post release monitoring programs are required not only to evaluate critically the degree of success and failure of biocontrol programs worldwide but also to provide insights into improving future biocontrol efforts. Handling Editor: Heikki Hokkanen.     

Plant-mediated interactions: Considerations for agent selection in weed biological control programs

Plant-mediated indirect interactions among herbivores (arthropods and pathogens) are common and extensively reported in the ecological literature. However, they are not well-documented with respect to weed biological control. Such interactions between biological control agents can have net positive or negative impacts on total weed suppression depending on the strength of the interaction(s), the relative importance of the agent indirectly impacted, and the combined weed suppression that results. A better understanding of plant-mediated interactions may improve decision-making about which agents to introduce in classical biological control programs for greatest impact on invasive weeds. This paper reviews the subject, including examples from the biological control literature; outlines the need for research on indirect effects of herbivores on other herbivores; discusses how such knowledge may strengthen classical biological control programs for invasive weeds; and provides recommendations for the kind of studies that should be done and how information about plant-mediated interactions could be integrated into agent evaluation protocols, to assist in decision-making about agents for importation and release.     

Melding ecology,classical weed biocontrol,and plant microbial ecology can inform improved practices in controlling invasive plant species

Early research leading to the successful biological control of invasive species such as Opuntia spp., and Hypericum perforatum set examples and provided data useful for research programs that would follow. However, this early work failed to become established as a source of applicable principles for later workers in weed biocontrol. Recently, retrospective and parallel studies have been suggested as a means to reengage with earlier work to derive useful ideas and data to enhance future programs in weed biocontrol. Parallel studies by workers in plant community ecology on the nature of feedback elicited by plant species in their invaded and native range have shown the importance of soil microbial communities in effecting feedback. Retrospective reexamination of previous studies would likely provide clues to other insect–plant pathogen interactions in addition to those described by the author and others. The effects of invasive species in profoundly altering soil microbial communities point to the need for further studies on key microbial species contributing to or driving the impact of biocontrol. These collective data suggest that the desired goal of selecting for and utilizing stronger biocontrol agents to reduce nontarget effects and to increase the impact of biological control programs would be best served by prerelease studies that assess the propensity of a candidate agent for direct or indirect interaction with other agents. This could be assessed through the use of survival analysis. Overall, parallel empirical and retrospective studies should be a necessary part of how biological control is practiced.     

Factors affecting the cost of weed biocontrol programs in New Zealand

Many national schemes for setting priorities for invasive weed management have emphasized the current or future impacts of the weed more than the cost or feasibility of control, perhaps because the latter may be difficult to estimate. As part of a project to improve prioritization of weed biocontrol targets in New Zealand, we investigated factors that were hypothesized to influence the cost of conducting weed biological control, using data from New Zealand programs. Taxonomic isolation of the target weed, relative to commercially important plants and native flora was not a significant influence on program cost, although we present evidence that disease, which to date has only affected agents released against taxonomically isolated weed targets, has masked the importance of taxonomic isolation in New Zealand. Opposition to biocontrol has caused delays, but has not had a major influence on the cost of biocontrol in New Zealand, probably because weed species with the greatest potential for opposition were identified during feasibility studies and avoided, or because conflicts were resolved by conducting cost-benefit analyses that were minor components of the total program costs. Only two factors explained virtually all the variance in program cost: program type (repeat programs were cheaper than novel/pioneering programs); and the number of agent species released. The predicted cost of future weed biocontrol programs can now be incorporated into decision-making tools ranking New Zealand weed biocontrol targets. Efficiencies in future programs are most likely to be gained by better agent selection so that fewer agents are released. For repeat programs this could be achieved by waiting until monitoring has been conducted overseas, so that the best agents or combination of agents can be selected for any particular weed. This reiterates the need for better post-release evaluation of weed biocontrol agent effectiveness worldwide.     

Tamarisk biocontrol,endangered species risk and resolution of conflict through riparian restoration

A long-standing debate between wildlife agencies and biological control researchers and practitioners concerns Diorhabda carinulata Desbrochers (Coleoptera: Chrysomelidae) introduced to suppress invasive Tamarix spp. (Tamaricaceae), and potential impacts of Tamarix defoliation on endangered southwestern willow flycatchers using this non-native plant as nesting habitat in some western riparian ecosystems. The conflict and ensuing legal actions are currently centered on the presence of D. carinulata within the breeding range of the flycatcher in the Virgin River watershed, which has led to APHIS termination of permits supporting the biocontrol development program and has also affected other programs to develop biocontrol agents against environmental weeds. Central to concerns over wildlife is the lack of rehabilitation of native vegetation where biocontrol is expected, so there are current and planned efforts to promote restoration of native cottonwood-willow habitat to mitigate the anticipation decline in Tamarix cover. A strategic approach to riparian restoration is outlined which could facilitate sustainable, and scientifically documented recovery of this iconic habitat type. While the results of these efforts will not be known immediately, the process which is leading to riparian restoration has brought specialists from both sides of the debate together in search of resolution via collaboration, and if successful, may allow re-initiation of the Tamarix biocontrol program attendant with habitat enhancement for wildlife species of conservation concern.     

Spatial and temporal variation in natural enemy assemblages on Maryland native plant species

Habitat manipulation is a branch of conservation biological control in which vegetation complexity and diversity are increased in managed landscapes to provide food and other resources for arthropod natural enemies. This is often achieved by maintaining noncrop plant material such as flowering strips and beetle banks that provide natural enemies with nectar and pollen, alternative prey, shelter from disturbance, and overwintering sites. In most cases, plant material used in habitat manipulation programs is not native to the area in which it is planted. Using native plant species in conservation biological control could serve a dual function of suppressing pest arthropod outbreaks and promoting other valuable ecosystem services associated with native plant communities. We evaluated 10 plant species native to Maryland for their attractiveness to foliar and ground-dwelling natural enemies. Plants that showed particular promise were Monarda punctata, Pycnanthemum tenuifolium, and Eupatorium hyssopifolium, which generally harbored the greatest abundance of foliar predators and parasitoids, although abundance varied over time. Among ground-dwelling natural enemies, total predator and parasitoid abundance differed between plant species, but carabid and spider abundance did not. Matching certain plant species and their allied natural enemies with specific pest complexes may be enhanced by identifying the composition of natural enemy assemblages at different times of year and in both foliar and ground habitat strata.     

Melding ecology, classical weed biocontrol, and plant microbial ecology can inform improved practices in controlling invasive plant species

Early research leading to the successful biological control of invasive species such as Opuntia spp., and Hypericum perforatum set examples and provided data useful for research programs that would follow. However, this early work failed to become established as a source of applicable principles for later workers in weed biocontrol. Recently, retrospective and parallel studies have been suggested as a means to reengage with earlier work to derive useful ideas and data to enhance future programs in weed biocontrol. Parallel studies by workers in plant community ecology on the nature of feedback elicited by plant species in their invaded and native range have shown the importance of soil microbial communities in effecting feedback. Retrospective reexamination of previous studies would likely provide clues to other insect–plant pathogen interactions in addition to those described by the author and others. The effects of invasive species in profoundly altering soil microbial communities point to the need for further studies on key microbial species contributing to or driving the impact of biocontrol. These collective data suggest that the desired goal of selecting for and utilizing stronger biocontrol agents to reduce nontarget effects and to increase the impact of biological control programs would be best served by prerelease studies that assess the propensity of a candidate agent for direct or indirect interaction with other agents. This could be assessed through the use of survival analysis. Overall, parallel empirical and retrospective studies should be a necessary part of how biological control is practiced.