Species distribution models for natural enemies of monarch butterfly (<Emphasis Type="Italic">Danaus plexippus</Emphasis>) larvae and pupae: distribution patterns and implications for conservation

Prey populations can be strongly influenced by predators and parasitoids, and migratory prey whose distributions vary geographically throughout their breeding seasons encounter different combinations of predators and parasitoids throughout their range. North American monarch butterflies (Danaus plexippus) are susceptible to a wide variety of natural enemies, but the distribution of these natural enemies has not been quantified. We developed ecological niche models using environmental data to identify areas with suitable abiotic conditions for eight known natural enemies of monarchs, including six predators: Arilus cristatus, Harmonia axyridis, Monomorium minimum, Podisus maculiventris, Polistes spp., and Solenopsis geminata; and two parasitoids: Lespesia archippivora and Pteromalus cassotis. We combined correlated suitable areas for individual predators and parasitoids to identify regions with the most predator and parasitoid species potential. The Gulf Coast, West Coast, Florida, and parts of the eastern United States are predicted to have the most natural enemy species. We suggest that future research should assess monarch mortality rates in these areas, and that monarch conservation strategies consider pressure from natural enemies.     

Biological control of the diamondback moth, Plutella xylostella: A review

The diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera: Plutellidae), is one of the most destructive cosmopolitan insect pests of brassicaceous crops. It was the first crop insect reported to be resistant to DDT and now, in many crucifer producing regions, it has shown significant resistance to almost every synthetic insecticide applied in the field. In certain parts of the world, economical production of crucifers has become almost impossible due to insecticidal control failures. Consequently, increased efforts worldwide have been undertaken to develop integrated pest management (IPM) programs, principally based on manipulation of its natural enemies. Although over 130 parasitoid species are known to attack various life stages of DBM, most control worldwide is achieved by relatively few hymenopteran species belonging to the ichneumonid genera Diadegma and Diadromus, the braconid genera Microplitis and Cotesia, and the eulophid genus Oomyzus. DBM populations native to different regions have genetic and biological differences, and specific parasitoid strains may be associated with the specific DBM strains. Therefore, accurate identification based on genetic studies of both host and parasitoid is of crucial importance to attaining successful control of DBM through inoculative or inundative releases. Although parasitoids of DBM larvae and pupae are currently its principal regulators, bacteria-derived products (e.g., crystal toxins from Bacillus thuringiensis) and myco-insecticides principally based on Zoophthora radicans and Beauveria bassiana are increasingly being applied or investigated for biological control. Viruses, nematodes and microsporidia also have potential as biopesticides for DBM. When an insect pest is exposed to more than one mortality factor, there is the possibility of interactions that can enhance, limit, or limit and enhance the various aspects of effectiveness of a particular control tactic. This paper reviews the effectiveness of various parasitoids and entomopathogens against DBM, interactions among them, and their possible integration into modern IPM programs.     

Biological studies on two neotropical egg parasitoid species: Trichogramma nerudai and Trichogramma sp. (Hymenoptera: Trichogrammatidae)

The main biological attributes of two Neotropical egg parasitoids, the arrhenotokous Trichogramma nerudai and the thelytokous Trichogramma sp., were assessed under controlled laboratory conditions. Developmental time from egg to adult, and parasitoid survival, fecundity and fertility were studied using life tables. Results showed that T. nerudai had a faster developmental time than Trichogramma sp. (13.014±0.4019 and 13.595±0.4931 days, respectively). Both species showed similar life table statistics, r_m was 0.222 and 0.225 for T. nerudai and Trichogramma sp., respectively. Parasitoid survival averaged 95% for both species. The results obtained are discussed in the context of selecting one of these natural enemies as a potential biological control agent for the European pine shoot moth Rhyacionia buoliana (Lepidoptera: Tortricidae) in pine forests and the codling moth Cydia pomonella (Lepidoptera: Tortricidae) in apple orchards in Argentina.     

Prey selection by Dicyphus hesperus of infected or parasitized greenhouse whitefly

The development of effective biological control programs in which predators are integrated with other natural enemies such as parasitoids and entomopathogens requires an understanding of their interactions. In this study we examined the extent to which the omnivorous mirid bug Dicyphus hesperus, an effective biological control agent of greenhouse whiteflies, accepts prey that are either parasitized by the specialist whitefly parasitoid, Encarsia formosa or infected by the generalist fungal entomopathogen, Beauveria bassiana. In non-choice laboratory experiments, we measured how parasitism and infection of greenhouse whitefly, Trialeurodes vaporariorum, as related to parasitoid age and course of fungal infection, might alter the probability of feeding by second instar or adult female predators. Predation by D. hesperus was similar for parasitized and unparasitized whiteflies, regardless of parasitoid age. However, predators generally avoided feeding on infected whiteflies, particularly when infection was manifested through the presence of oosporein or hyphae on the surface of prey.     

Tea: Biological control of insect and mite pests in China

Tea is one of the most economically important crops in China. To secure its production and quality, biological control measures within the context of integrated pest management (IPM) has been widely popularized in China. IMP programs also provide better control of arthropod pests on tea with less chemical insecticide usage and minimal impact on the environment. More than 1100 species of natural enemies including about 80 species of viruses, 40 species of fungi, 240 species of parasitoids and 600 species of predators, as well as several species of bacteria have been recorded in tea ecosystems in China. Biological and ecological characteristics of some dominant natural enemies have been well documented. Several viral, bacterial, and fungal insecticides have been commercially utilized at large scale in China. Progress in biological control methods in conjunction with other pest control approaches for tea insect pest management is reviewed in this article. Knowledge gaps and future directions for tea pest management are also discussed.     

12种杀虫剂及5种天敌昆虫对梧桐木虱的田间控害评价

梧桐木虱是一种危害园林植物梧桐的重大害虫。本研究分别测定了12种杀虫剂和5种天敌昆虫对梧桐木虱的防治效果。结果表明: 12种杀虫剂中噻虫嗪和吡虫啉对梧桐木虱的防治效果最好,螺虫乙酯次之。在喷雾法、注射法和灌根法等3种施药方式中,应尽量避免使用喷雾法,优先选择注射法,在条件允许的情况下可使用灌根法作为补充手段。5种天敌昆虫中以中华通草蛉和赤星瓢虫对梧桐木虱的防治效果最佳,在长期防治效果上可以起到代替杀虫剂的作用。因此,中华通草蛉及赤星瓢虫可作为园林环境中长期安全环保防治梧桐木虱的重要方法,而以注射法使用噻虫嗪和吡虫啉可作为短期快速防治手段。     

Is the Naturally Derived Insecticide Spinosad® Compatible with Insect Natural Enemies?

Spinosad® (Dow Agrosciences) is a neurotoxic insecticide produced by fermentation of an actinomycete. Spinosad is classified as an environmentally and toxicologically reduced risk material and has been embraced by IPM practitioners as a biorational pesticide. We examined the available information on the impact of spinosad on natural enemies and classified mortality responses to spinosad using the IOBC laboratory and field scales that run from 1 (harmless) to 4 (harmful). In total, there were 228 observations on 52 species of natural enemies, of which 162 involved predators (27 species) and 66 involved parasitoids (25 species). Overall, 71% (42/59) of laboratory studies and 79% (81/103) of field-type studies on predators gave a class 1 result (not harmful). Hymenopteran parasitoids are significantly more susceptible to spinosad than predatory insects with 78% (35/45) of laboratory studies and 86% (18/21) of field-type studies returning a moderately harmful or harmful result. Predators generally suffer insignificant sub-lethal effects following exposure to spinosad, whereas parasitoids often show sub-lethal effects including loss of reproductive capacity, reduced longevity, etc. All studies agree that spinosad residues degrade quickly in the field, with little residual toxicity at 3-7 days post-application. We also examined the importance of route of exposure, species-specific and stage-specific susceptibility and we make recommendations for future studies. We conclude that for conservation of predator populations, spinosad represents one of the most judicious insecticides available but the use of this product should be evaluated carefully in situations where conservation of parasitoid populations is of prime concern.     

Biological control of aphids and coccids: a comparative analysis

A comparison of the biological control of aphids and coccids was carried out by analyzing success rates for the three major types of biological control, i.e., classical, augmentative, and conservational. Because of the higher intrinsic rates of increase for aphids versus coccids, the working hypothesis that biological control of aphids is less successful compared to coccids was adopted. However, this hypothesis was not supported by an analysis of classical biological control using the BIOCAT database. In this analysis, parasitoids were more successful than predators when used against either aphids or coccids, but the control of Icerya spp. with Rodolia spp. (predators) was highly successful. Some reasons for success of Rodolia spp. are adduced, but field studies on the long-term population dynamics of Icerya–Rodolia systems are needed for determining the mechanisms of regulation. Comparative analyses of augmentative and conservational biological control of aphids and coccids were inconclusive, due to lack of adequate databases; some possible factors involved in the success of these types of biological control are discussed. It is suggested that parasitoids could be better control agents than predators in augmentative biological control of aphids in production greenhouses. Conservational biological control of either aphids or coccids should be aimed at enhancing populations of indigenous natural enemies, especially mobile generalist predators that are capable of keeping pace with mobile pests.     

Development and application of an integrated pest management programme for the psyllid,Ctenarytaina thysanura onBoronia megastigma in Tasmania

An integrated pest management programme for control of the psyllid,Ctenarytaina thysanura (Ferris & Klyver) (Hemiptera: Psyllidae) in commercial boronia plantations was developed and implemented from 1986–1989. The programme involved spraying boronia stems only with the systemic organophosphate mevinphos at peak incidence of late stage nymphs. At this time, the majority of parasitoids were pupating within mummified hosts. Consequently, actively feeding nymphs were potentially susceptible to mevinphos but the parasitoids were not. Stem application negated any contact effect on parasitoids and predators. The conservation of natural enemies subsequently reduced psyllid numbers and no insecticide has been required since 1989. In economic terms the benefit to cost ratio of the IPM programme was $22.40: $1.0 and this was 40 times greater than the conventional 10 demeton-S-methyl sprays used by growers prior to the study.     

Effect of flower traits and hosts on the abundance of parasitoids in perennial multiple species wildflower strips sown within oilseed rape (<Emphasis Type="Italic">Brassica napus</Emphasis>) crops

Reducing the use of insecticides is an important issue for agriculture today. Sowing wildflower strips along field margins or within crops represents a promising tool to support natural enemy populations in agricultural landscapes and, thus, enhance conservation biological control. However, it is important to sow appropriate flower species that attract natural enemies efficiently. The presence of prey and hosts may also guide natural enemies to wildflower strips, potentially preventing them from migrating into adjacent crops. Here, we assessed how seven flower traits, along with the abundance of pollen beetles (Meligethes spp., Coleoptera: Nitidulidae) and true weevils (Ceutorhynchus spp., Coleoptera: Curculionidae), affect the density of parasitoids of these two coleopterans in wildflower strips sown in an oilseed rape field in Gembloux (Belgium). Only flower traits, not host (i.e. pollen beetles and true weevils) abundance, significantly affected the density of parasitoids. Flower colour, ultraviolet reflectance and nectar availability were the main drivers affecting parasitoids. These results demonstrate how parasitoids of oilseed rape pests react to flower cues under field conditions. Similar analyses on the pests and natural enemies of other crops are expected to help to develop perennial flower mixtures able to enhance biological control throughout a rotation system.     

Current status and future perspectives on natural enemies for pest control in Taiwan

ABSTRACT

In Taiwan, the agricultural policy, ‘Reduce the consumption of pesticide to half in the next 10 years’, was launched in 2017. Pesticide application, which results in contamination of food by chemical residues, pest resistance, and other adverse ecological effects, is a growing public and environmental concern. Pest control by natural predators is, thus, the best alternative. Biological control methods implemented based on insights obtained from studies on pest behaviour, rearing, and various crop management modes, increase the possibility of controlling pests in modern organic agricultural systems. More than a decade has passed since the first introduction of a predatory insect in Taiwan for pest control (in the 1990s). Predatory and parasitic natural enemies, including lacewing, predatory stink bugs, Orius, and parasitic wasps, were initially used for controlling thrips, aphids, spider mites, whiteflies, and lepidopteran pests. At present, there exists a wide range of integrated pest management (IPM) methods incorporating other non-chemical, biological, and agricultural methods. However, recently, there has been an increase in research and development on the utilisation of natural enemies of insects and the associated food safety issues. Mass production and release, storage, and handling techniques of insect predators and parasitoids have been successful in recent years. The final goal of present day research is to develop natural enemy products and provide an IPM-based model to farmers for using natural enemies in agricultural production systems, thereby reducing pesticide application and ensuring food security.     

Preface

Data on the toxicity and selectivity of synthetic pyrethroids (SPs) to arthropod natural enemies and their host or prey are reviewed with emphasis on cotton, apple, alfalfa, cereal and vegetable inhabiting species. Generally, SPs are variably toxic and selective (in relation to their hosts or prey) to species within most families of natural enemies. Exceptions are low to moderate toxicity and favorable selectivity to most hemipteran predators, and high toxicity and unfavorable selectivity to virtually all phytoseiid mites in comparison to their prey. In North America, SPs are more favorably selective to cotton natural enemies over their prey or host than to apple inhabiting species. These differences may be due to intrinsic levels of susceptibility (preselection levels) between the types of natural enemies exploited on both crops in IPM programs (i.e. hemipteran species on cotton versus phytoseiid mites on apple) and/or to tolerances or resistances differences due to previous patterns of chemical use on these crops. Possible means of increasing selective use of SPs in future IPM systems based on ecological and physiological selectivity including the development of SP-resistant predators are discussed.     

Natural enemies and environmental factors affecting the population dynamics of the gypsy moth

The population densities of the gypsy moth (Lymantria dispar; Lepidoptera: Lymantriidae) may reach outbreak levels that pose considerable economic and environmental impacts to forests in Europe, Asia, Africa and North America. Compared with the situation in its native European range feeding damage by gypsy moth is often found to be more severe in North America and other parts of the world. Thus, the release from natural enemies can be interpreted as an important cause for high feeding damages. Natural enemies, especially parasitoids, can cause delayed density‐dependent mortality, which may be responsible for population cycles. In North America where only few parasitoids have been introduced and the parasitism rates are considerably lower than in Europe, generalist predators play a larger role than in Europe. Many other factors seem to influence the population dynamics of the gypsy moth such as the host plants and weather. Nevertheless, much of the variability in population densities of the gypsy moth may be attributed to interacting effects of weather conditions and attack by natural enemies. In spite of the considerable number of studies on the ecology and population dynamics of the gypsy moth and the impact of their natural enemies, more quantitative information is required to predict the population dynamics of this pest species and to control its economic and ecologic impact.     

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.     

Prospects for the biological control of subterranean termites (Isoptera: rhinotermitidae), with special reference to Coptotermes formosanus

Costs associated with subterranean termite damage and control are estimated to approach $2 billion annually in the United States alone. The Formosan subterranean termite, Coptotermes formosanus Shiraki, is one of the more economically important subterranean species. In recent years, the shortcomings associated with conventional chemical control methods have prompted policymakers and scientists to evaluate the potential for biological control of subterranean termites (C. formosanus in particular), that is, to determine the potential for natural enemies - predators, parasitoids and pathogens - to suppress termite populations. Ants are the greatest predators of termites, and may have a considerable local impact on termite populations in some areas of the world. A few parasitoids of termites are known, but their potential for regulating termite populations seems negligible. Characteristics of the colony, such as a protected, underground location (and, for the C. formosanus nest, its modular and dispersed nature), are likely to limit the impact predators and parasitoids have on subterranean termites. Thus, there seems little potential for use of these agents for subterranean termite control. For various reasons, pathogenic organisms, such as viruses, bacteria, Protozoa, nematodes and most fungi, have shown little promise for use in biological termite control. However, research suggests that strains of two well-studied, endoparasitic fungi, Beauveria bassiana and Metarhizium anisopliae, when employed in baiting schemes, may offer the potential for at least some measure of subterranean termite control, although their successful use is compromised by a number of inherent biological limitations and logistical problems that have yet to be solved. Although not strictly in the realm of classical biological control, recent studies suggest that natural products, such as ant semiochemicals and fungal metabolites (siderophores), or their synthetic analogues, eventually might find a use in termite control programmes as repellents or insecticides in wood treatments or soil applications if stable formulations can be developed.     

Arthropod Natural Enemies of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) in India

Helicoverpa armigera Hubner Lepidoptera: Noctuidae is one of the most serious insect pests in the Old World. In India, it causes substantial losses to legume, fibre, cereal oilseed and vegetable crops. This paper reviews the literature on the biology, ecology, efficacy, rearing and augmentation of endemic parasitoids and predators, as well as exotic parasitoids introduced and released in India. It also provides updated lists of H. armigera natural enemies native to India. In addition, reports of augmentative releases of Trichogramma spp., the most extensively studied natural enemy of H. armigera are summarized.     

Trichoderma spp. as elicitors of wheat plant defense responses against Septoria tritici

Leaf blotch of wheat is a widespread and highly active disease that affects wheat production. In addition to the use of chemicals and proper cultivation methods, microbial antagonists are used to control plant pathogens. Trichoderma spp. stimulate a systemic induced response in plants. Therefore, the efficacy of Trichoderma spp. against wheat leaf blotch was evaluated under greenhouse conditions. The susceptible plants were sprayed with Septoria tritici conidiospores. In order to select an efficient method of pretreatment with Trichoderma spp., leaf spraying and seed coating with 14 isolates were tested in 2003 and 2004. The extent of leaf necrosis area and pycnidial coverage was estimated. Antagonism was assessed by the capacity of each Trichoderma spp. isolate to restrict the progress of leaf blotch, 21 days after inoculation. Of the two methods, seed coating was more efficacious against leaf blotch than leaf spraying. Amongst the 14 isolates tested, the isolate prepared from T. harzianum (Th5) produced the highest level of protection. None of the treatments caused changes in plant stem diameter or dry weight. Trichoderma spp. did not get into leaves while S. tritici was present, even in asymptomatic leaf extracts. In addition, the leaf apoplast antifungal proteolytic activity was measured in plants 7, 15, and 22 days after sowing. This antifungal action decreased in plants only inoculated with S. tritici, but increased in those grown from seeds coated with the T. harzianum (Th5) isolate. This increase conferred resistance to the susceptible wheat cultivar. The endogenous germin-like protease inhibitor coordinated the proteolytic action. These results suggest that T. harzianum stimulates a biochemical systemic induced response against leaf blotch.     

Combining lacewings and parasitoids for biological control of foxglove aphids in sweet pepper

The role of natural enemy diversity in biological pest control has been debated in many studies, and understanding how interactions amongst predators and parasitoids affect herbivore populations is crucial for pest management. In this study, we assessed the individual and combined use of two species of natural enemies, the parasitoid Aphidius ervi Haliday, and the predatory brown lacewing Micromus variegatus (Fabricius), on their shared prey, the foxglove aphid, Aulacorthum solani (Kaltenbach), on sweet pepper. We hypothesized that the presence of intraguild predation (IGP) and predator facilitation (through induced aphid dropping behaviour) might have both negative and positive effects on aphid control, respectively. Our greenhouse trial showed that overall, the greatest suppression of aphids occurred in the treatment with both the parasitoid and the lacewing. While the combination of lacewings and parasitoids significantly increased aphid control compared to the use of parasitoids alone, the effect was not significantly different to the treatment with only predators, although there was a clear trend of enhanced suppression. Thus, the combined effects of both species of natural enemies were between additive and non‐additive, suggesting that the combination is neither positive nor negative for aphid control. High levels of IGP, as proven in the laboratory, were probably compensated for by the strong aphid suppression provided by the lacewings, whether or not supplemented with some level of predator facilitation. For aphid management over a longer time scale, it might still be useful to combine lacewings and parasitoids to ensure stable and resilient aphid control.     

Rise and fall of cotton aphid (Hemiptera: Aphididae) populations in southeastern cotton production systems

The impact of natural enemies on cotton aphid, Aphis gossypii Glover (Hemiptera: Aphididae), populations in cotton, Gossypium hirsutum L., production systems in the southeastern United States was evaluated over 3 yr in irrigated commercial cotton fields. Fungal epizootics caused by the entomopathogen Neozygites fresenii (Nowakowski) Batko reduced aphid numbers to subthreshold levels in 1999, 2000, and 2001 and occurred consistently in early to mid-July in all 3 yr. Scymnus spp. were the most abundant aphidophagous predators, although other coccinellid species and generalist predators such as spiders, fire ants, heteropterans, and neuropterans also were present. Studies using arthropod exclusion cages demonstrated little impact of predators or parasitoids on aphid populations before fungal epizootics. Arthropod natural enemies were most abundant after epizootics and may have suppressed aphid populations late in the season. Seed cotton yield, and lint quality were not affected by aphicide applications in any year of the study. Implications of these findings for aphid management in the southeastern United States are discussed.     

Evolutionary ecology of the interactions between aphids and their parasitoids

Many organisms, including entomopathogenous fungi, predators or parasites, use aphids as ressources. Parasites of aphids are mostly endoparasitoid insects, i.e. insects which lay eggs inside the body of an other insect which will die as a result of their development. In this article, we review the consequences of the numerous pecularities of aphid biology and ecology for their endoparasitoids, notably the Aphidiinae (Hymenoptera: Braconidae). We first examine the various mechanisms used by aphids for defence against these enemies. We then explore the strategies used by aphidiine parasitoids to exploit their aphid hosts. Finally, we consider the responses of both aphids and parasitoids to ecological constraints induced by seasonal cycles and to environmental variations linked to host plants and climate. The fundamental and applied interest of studying these organisms is discussed.