Landscape context and management effects on an important insect pest and its natural enemies in almond

Pest control mediated by organisms such as parasitoids is a valuable ecosystem service, particularly with regard to high costs, low effectiveness, and detrimental effects of some agrochemicals. This study examined infestation rates and abundance of pests and their natural enemies in organic and conventional almond orchards in California, differing in landscape context, understory plant cover, and plant species richness. Parasitoids of the commercially most important insect pest of almond, the Navel Orangeworm (NOW) were studied by rearing NOW in collected overwintering nuts. The indirect impact of vertebrate natural enemies of NOW were estimated by counting empty nut shells with feeding marks by wild birds and various mammals, found at the orchard floor. Mean nut infestation by NOW ranged from 0.8% to 37% per orchard and was reduced by parasitism rates, ranging from 0% to 22%, and vertebrate nut damage, ranging from 2% to 96% per orchard. The parasitoids were facilitated by a high proportion of natural habitat surrounding the orchards and high proportion of understory ground cover with vegetation. The vertebrate natural enemies were facilitated by a high proportion of natural habitat surrounding the orchards and plant species richness in the orchard understory. In conclusion, this study shows that pest control mediated by vertebrates and invertebrates promoted by near natural habitats can lower pest pressure by NOW larvae in overwintering almond. In case of the vertebrate nut damage this service might only be temporal and turn into a dis-service during and after harvest because the vertebrates continue to feed on the nuts and may also cause injuries to the trees.     

Biocontrol of Pests of Apples and Pears in Northern and Central Europe: 2. Parasitoids

The parasitoids of arthropod pests of apple and pear in northern and central Europe and their use as biological control agents are reviewed. The review demonstrates that apple and pear pests are host to a large and varied parasitoid fauna. All important pests are known to be host of parasitoids, but many parasitoids play only a minor part in regulating populations of their host. However, many parasitoid species are important natural enemies and some effectively regulate pest populations in unsprayed and/or commercial (insecticide sprayed) apple or pear orchards either individually or as part of parasitoid guilds. Exploitation/fostering of existing populations of parasitoids has been demonstrated to be an effective or partially effective approach for natural control of several important pest species. Important examples include natural regulation of the apple sawfly by Lathrolestes ensator and Aptesis nigrocincta, of the summer fruit tortrix moth by Colpoclypeus florus and Teleutaea striata, of leaf midges by Platygaster demades, of woolly aphid by Aphelinus mali and of leaf mining moths by guilds of parasitoid species. Introduction of parasitoids is an alternative approach to the exploitation of parasitoids already present in the orchard. This approach has been little explored and its success rate has been low, mainly confined to the control of non-indigenous pests by introducing parasitoids from their native region. Mass production methods for parasitoids are difficult and costly and are likely to be economic only where long-term populations can be established. Even where low cost mass culture techniques are developed, the degree of control may not be high enough to prevent economic pest damage as demonstrated by negative results with mass release of Trichogramma egg parasites for control of tortricids in orchards. Suitability of the orchard habitat is recognized as crucial to the success of individual parasitoids. Key requirements are adequate populations of the pest(s) and/or alternative hosts, suitable shelter, overwintering sites or food sources and avoidance of harmful effects of pesticides. Many species are highly sensitive to broad-spectrum insecticides, especially in the adult life-stage. Avoiding the harmful affects of insecticides is crucial to successful exploitation. The use of insecticides needs to be avoided, either altogether or at crucial times in the parasitoids' life cycle, or less harmful alternatives need to be used. Numerous parasitoids could potentially be exploited as biological control agents but hitherto have received little attention because little is known about them and/or because they are sensitive to broad-spectrum pesticides and are thus virtually absent from commercial orchards. The aim of future studies should be to develop effective strategies for establishing equilibria between important pests and their parasitoids, with pest damage rarely exceeding the economic threshold.     

Seasonal biology and abundance of Psyllaephagus pistaciae,a biocontrol agent of the common pistachio psylla Agonoscena pistaciae

The solitary endoparasitoid Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), is the most widely distributed biological control agent of the common pistachio psyllid, Agonoscena pistaciae Burckhardt and Lauterer (Hemiptera: Psylloidea), in Iran. The pupation and overwintering sites of diapausing parasitoids and the psyllid were studied for 2 years using emergence traps in pistachio orchards in Rafsanjan, Iran. The psyllid mummies containing the overwintering parasitoid adhered to pistachio leaves and were carried on these leaves away from the tree when they latter senesced. The present results verified that plant litter which included dried grasses and old pistachio leaves tended to support a greater population of adult winter-form psyllid and psyllid mummies during the winter through early spring than other options examined. Adult parasitoids appeared in the field in early April, about 30 days after the emergence of adult psyllids, but almost at the same time as the hatching of the first generation psyllid nymphs in early April. Rates of parasitism of CPP were generally low throughout most of the year, ranging from 1 to 5%, but rose in late autumn to about 11%. Results suggest that the density of P. pistaciae is not great enough to keep pace with the psyllid populations in these orchards. They explain why growers consider it necessary to apply pesticides for this pest. However, this parasitoid undoubtedly does play an important role in the natural control of A. pistaciae late in the growing season, particularly in non-sprayed orchards. Conservation of these natural enemies should be one of the objectives in the development of sustainable pest management programs.     

Can the Understory Affect the Hymenoptera Parasitoids in a Eucalyptus Plantation?

The understory in forest plantations can increase richness and diversity of natural enemies due to greater plant species richness. The objective of this study was to test the hypothesis that the presence of the understory and climatic season in the region (wet or dry) can increase the richness and abundance of Hymenoptera parasitoids in Eucalyptus plantations, in the municipality of Belo Oriente, Minas Gerais State, Brazil. In each eucalyptus cultivation (five areas of cultivation) ten Malaise traps were installed, five with the understory and five without it. A total of 9,639 individuals from 30 families of the Hymenoptera parasitoids were collected, with Mymaridae, Scelionidae, Encyrtidae and Braconidae being the most collected ones with 4,934, 1,212, 619 and 612 individuals, respectively. The eucalyptus stands with and without the understory showed percentage of individuals 45.65% and 54.35% collected, respectively. The understory did not represent a positive effect on the overall abundance of the individuals Hymenoptera in the E. grandis stands, but rather exerted a positive effect on the specific families of the parasitoids of this order.     

How many species are there in apple insect communities?: testing the resource diversity and intermediate disturbance hypotheses

Abstract.

• 1 In European and American apple orchards the insect species richness, calculated from our own, and published data, varied widely (30–940 species). The dominance of insect orders was similar to that found in natural communities.
• 2 To test the predictions of the‘resource diversity hypothesis’(RDH) and the‘intermediate disturbance hypothesis’(IDH) of insect diversity, we analysed the simultaneous impact of vegetational diversity and intensity of orchard management practice on the species richness of characteristic insect categories in six types of apple stands, over 5 years, in Hungary.
• 3 The more diverse was the vegetation adjacent to the orchard and the less intensively was the orchard managed, the greater was the total insect species richness, supporting both hypotheses.
• 4 The number of agricultural pest species found in the apple orchards depended above all on the diversity of adjacent agricultural vegetation, supporting only the RDH.
• 5 The species richness of specialized apple pests was limited only by the intensity of pest control, supporting only the IDH.
• 6 Within the category of generalist natural enemies species richness mainly varied with the diversity of adjacent vegetation, supporting the RDH.
• 7 The aeroentomofauna constituted a surprisingly high percentage (>50%) for both total species and agricultural pests, but was poorly represented within the natural enemies category, and absent from the specific apple pest category.
• 8 A general conclusion is that the number of insect species in perennial orchard-systems within agricultural areas is determined primarily by the regional extra-orchard vegetational diversity and secondly by the degree of local intra-orchard disturbances and plant diversity. Consequently, at the community level the predictions of both hypotheses can be simultaneously valid at different spatial scales and are not mutually exclusive. However, within the community, at the level of the different insect categories studied here, only one of the hypotheses proved to be valid.

     

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.     

More intraguild prey than pest species in arachnid diets may compromise biological control in apple orchards

Understanding the full diet of natural enemies is necessary for evaluating their role as biocontrol agents, because many enemy species do not only feed on pests but also on other natural enemies. Such intraguild predation can compromise pest control if the consumed enemies are actually better for pest control than their predators. In this study, we used gut metabarcoding to quantify diets of all common arachnid species in Swedish and Spanish apple orchards. For this purpose, we designed new primers that reduce amplification of arachnid predators while retaining high amplification of all prey groups. Results suggest that most arachnids consume a large range of putative pest species on apple but also a high proportion of other natural enemies, where the latter constitute almost a third of all prey sequences. Intraguild predation also varied between regions, with a larger content of heteropteran bugs in arachnid guts from Spanish orchards, but not between orchard types. There was also a tendency for cursorial spiders to have more intraguild prey in the gut than web spiders. Two groups that may be overlooked as important biocontrol agents in apple orchards seem to be theridiid web spiders and opilionids, where the latter had several small-bodied pest species in the gut. These results thus provide important guidance for what arachnid groups should be targets of management actions, even though additional information is needed to quantify all direct and indirect interactions occurring in the complex arthropod food webs in fruit orchards.     

Influence of the margin vegetation on the conservation of aphid biological control in apple orchards

The influence of three margin strip treatments (wildflower strips, grass strips and spontaneous vegetation) adjacent to apple orchards on the biological control of Dysaphis plantaginea Passerini (Hemiptera: Aphididae) was compared during two consecutive years. The wildflower strips provided the highest amount of floral resources. Within the margin strips, hoverflies responded positively to higher resource provisioning whereas ladybird abundance did not differ between strip treatments. Within the orchards, the presence of parasitoids, hoverflies, and ladybirds in aphid colonies and the predation of sentinel aphids were not significantly affected by the adjacent strip treatments. The number of natural enemies observed in aphid colonies was mainly driven by aphid number. Aphid numbers were higher close to the margin strips suggesting that aphid colonization from orchard edges may counteract the positive effect of wildflower strips on natural enemy abundance and on a reduction of aphid infestation. The results confirm the positive influence of floral resource provisioning by wildflower strips on the conservation of aphid natural enemies, but also suggest that effects of wildflower strips on aphid regulation inside orchards are not very strong compared with spontaneous vegetation naturally occurring in the margins.     

Life cycle,phenology and economic importance of the walnut husk fly Rhagoletis completa Cresson (Diptera: Tephritidae) in northern Italy

The walnut husk fly Rhagoletis completa Cresson, a pest originating from North-America, was detected for the first time in Europe in 1991. The life cycle and phenology of R. completa were studied, during two successive seasons, in two commercial orchards located in northern Italy. The pest develops one generation per year. Fly emergence lasted from early July to the second half of August. Oviposition was detected from late July to early September, with peaks between August 5 and 18. First instar larvae were recorded from early August and mature larvae left husks from late August onwards to pupate in the soil. The seasonal patterns of nuts infestation showed rapid growths in August following oviposition peaks. Nuts infestation levels in the untreated plots of the two orchards, ranged from 74–91% in the first year to 89–91% in the following season. Relatively dark shell surfaces were linked to a weight reduction in nuts and kernels and to an increase in darkened and mouldy kernels. Observations made in an untreated orchard for additional 3 years confirmed these trends. Baited Pherocon AM were effective in monitoring flies but catches on woody green spheres were better correlated with the first relevant oviposition phases. When pesticides were sprayed with the correct timing, i.e. against eggs or first instar larvae, infestation was kept to acceptable levels with a single application. Considering average yields, nuts prices and costs for insecticide use, 1–2 insecticide treatments are economically viable. Other Implications are discussed.     

Biological control of insect pests in apple orchards in China

Apple is one of the most important fruits in China, and both yield and quality are greatly affected by insect pests. According to surveys, there are more than 200 species of natural enemies in apple orchards. Few, however, have been closely studied. Major natural enemies including parasitoids, predators and pathogens are briefly described in this review, especially focusing on two parasitoids of Trichogramma dendrolimi Matsumura and Aphelinus mali Haldeman, predatory mites and a pathogenic fungus of Beauveria bassiana (Balsamo) Vuillemin as case studies. Augmentation, one important strategy of biological control, supplements the natural control provided by the existing natural enemy community in apple orchards, and greatly increases their efficiency in controlling pests. Conservation biological control is also widely applied in four major apple-producing areas. Based on habitat manipulation, the ground cover planting system helps regulate the microclimate and enhance the biodiversity of apple orchards, effectively conserving the richness and diversity of beneficial insect species. Certain achievements have been made in the main biological control strategies including successful introduction of some exotic natural enemies such as A. mali and Typhlodromus occidentalis Nesbitt, augmentative production and application of biological control agents such as T. dendrolimi, B. bassiana and Bacillus thuringiensis, and further research in conservation of establishing adaptive ground cover planting patterns to local environment. Challenges, however, still exist. Biological control of insect pests in apple orchards is an important part of integrated pest management programs, requiring more research and application in China.     

Diversity of natural enemies in Central Otago,New Zealand apple orchards: a practical measure of sustainability in pest management?

The diversity of the arthropod fauna in apple orchards in Central Otago was recorded from 1994 to 1999 using beating trays, pitfall traps, and sticky traps and the data analysed by the Shannon–Wiener Index. Three different fruit production systems were compared, conventional (CFP), integrated (IFP), and biological (BFP), to determine whether total arthropod diversity and/or the diversity of natural enemies (predators and parasitoids) could be used to measure the sustainability of their pest management practices. The contribution of natural enemies to the diversity indices was also examined to determine if they reflected the key species essential for sustainable pest management. The diversity of natural enemies in beating trays was found to be the measure most sensitive to changes in pest management. The diversity of both the total arthropod fauna and natural enemies was much lower under CFP than under either IFP or BFP, due to the use of broad-spectrum pesticides. This practice made the CFP programme unsustainable and it is no longer used. During the transition from CFP to IFP, the diversity indices rose to become similar to that in the BFP programme by the third season. The introduction of frequent applications of fungicides in BFP, including lime sulphur, appeared to reduce total arthropod and natural enemy diversity. Falls in the diversity indices for natural enemies were shown to reflect reduced numbers, range, and evenness of natural enemies but not necessarily the key species known to be critical for sustainable pest management. It is proposed that a Shannon–Wiener Index for natural enemies in beating trays of 0.2 or less in summer is strongly indicative of unsustainable pest management in Otago apple orchards. On the other hand, indices of 0.3–0.8, as found under IFP and BFP, do not give unequivocal or quantitative indications of the sustainability of pest management. A suite of other measures of sustainability are more useful, notably combining lower pesticide use, reduced pest damage and presence at harvest, the greater roles of key natural enemies, more effective plant resistance to pests and diseases, and higher profitability.     

Relationships between management practices and ground‐active invertebrate biodiversity in New Zealand kiwifruit orchards

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Entomofauna associated to horticultural crops under organic and conventional practices in Córdoba, Argentina

Farming practices and the addition of chemical synthetic substances in conventional agroecosystems are detrimental mainly to natural enemies of phytophagous insects, diminishing the natural regulation of pest insects. On the other hand, in organic agriculture, biological processes and care of the environment are favoured, hence an increase in insect biodiversity is predicted in this type of systems. In this work, abundance, richness of insects and proportion of functional groups were compared through a single quantitative sampling of insects in horticultural crop fields, three under organic and three under conventional management practices. Insect species richness, total and for guilds (phytophagous and entomophagous insects) were significantly higher in organic orchards, and also was the abundance of entomophagous insects. Richness and abundance of all insect orders (with exception of Homoptera abundance), were higher in orchards under organic management, being significative the differences for richness of Coleoptera and richness and abundance of Hymenoptera. Similar tendencies were observed in data obtained through sweep net in weeds. These results suggest that organic practices increase the diversity of species, particularly that of natural enemies.     

间种扁茎黄芪牧草枣园害虫天敌功能团的组成与时空动态

为了有效地管理枣树害虫,2002年在山西省太谷地区对4种不同处理的间作牧草枣园天敌功能团的群落组成、数量时空和多样性的发生动态进行了系统的调查研究.结果表明,间种牧草枣园害虫的天敌明显大于未间种牧草的枣园,种草综合防治园、种草常规防治园和种草不防治园的天敌比未种草常规防治园分别增加187.99%、151.82%和210.03%.种草枣园天敌功能团的恢复与重建速率、时空二维生态位平均宽度与重叠程度,以及平均多样性指数均明显大于未种草枣园;而种草枣园多样性标准误的波动幅度明显小于未种草枣园;不同处理枣园的天敌功能团与物种的优势度、多样性和均匀度的变化趋势均极显著相关.说明枣园种草有利于保护和利用天敌,有利于天敌的扩繁增殖;种草枣园天敌群落比未种草枣园相对稳定;不同时期、不同处理枣园天敌功能团数量和所起的作用不同;在天敌功能团中,瓢虫类恢复能力较强,蜘蛛类与寄生性天敌类较弱.用功能团来代替物种进行枣园害虫天敌群落学研究是可行的.     

Flower strips increase the control of rosy apple aphids after parasitoid releases in an apple orchard

Mass releases of two parasitoid species, Aphidius matricariae and Ephedrus cerasicola, may provide an alternative measure to pesticides to control the rosy apple aphid Dysaphis plantaginea in organic apple orchards. As an exploratory study, we tested if the presence of flower strips between apple tree rows could improve the action of three early parasitoid releases––and of other naturally present aphid enemies––on the control of aphid colonies and the number of aphids per tree. Apple trees located at various distances from parasitoid release points were monitored in plots with and without flower strips in an organic apple orchard over two years, along the season of aphid infestation (March to July). Our case study demonstrated that the presence of flowering plant mixes in the alleyways of the apple orchard reduced the presence of D. plantaginea by 33.4%, compared to plots without flower strips, at the infestation peak date. We also showed a negative effect of increasing the distance to parasitoid release points on aphid control. However, our results at the infestation peak date suggest that the presence of flower strips could marginally compensate for the detrimental effect of increasing distance to the release point, probably by improving the persistence and dispersal capacities of natural enemies. Despite high variations in aphid population dynamics between years, we conclude that combining flower strips with early parasitoid releases in apple orchards is promising for biological control of the rosy apple aphid, although the method merits to be further refined.     

Do weaver ants affect arthropod diversity and the natural‐enemy‐to‐pest ratio in horticultural systems?

High biodiversity is an important component of sustainable agricultural systems, and previous studies have found that increases in the diversity of the natural enemies of pests are associated with decreases in pest populations. Weaver ants are well known for their highly territorial and aggressive behaviour and for their control efficiency of many insect pests in tropical crop trees. Because of this, the ants have been used as a key component in integrated pest management (IPM) programmes for tropical crop trees. In implementing the IPM programmes, we received a number of enquiries related to whether weaver ants have negative effects on arthropod diversity and other natural enemies in orchard systems due to their aggressive behaviour. To answer these questions, we regularly sampled canopy arthropods in cashew and mango orchards in the Northern Territory of Australia in 1996, 2002 and 2003. We sampled, using a vacuum sampler, orchards with and without weaver ants. Cashew and mango plots with abundant weaver ants had similar or higher canopy arthropod and natural enemy diversity and similar ratios of natural enemies to insect pests, compared with plot where the weaver ant was absent. The study also showed that the application of insecticides reduced arthropod diversity and the ratio of natural enemies to insect pests in a mango orchard. However, insecticide spray did not affect natural enemy diversity and abundance, which may be related to a high immigration rate of natural enemies in small plots surrounded by areas that were not sprayed.     

Evaluation of monitoring traps with novel bait for navel orangeworm (Lepidoptera: Pyralidae) in California almond and pistachio orchards

Experiments conducted in three almond, Prunus dulcis (Rosales: Rosaceae), orchards and three pistachio, Pistacia vera (Sapindales: Anicardiaceae), orchards in 2009 and 2010, and determined that sticky bottom wing traps baited with ground pistachio mummies, or a combination of ground pistachio plus ground almond mummies, trapped more adult female navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae), than did traps baited with ground almond mummies alone. During both years of this study, 2.9 and 1.8 more moths were caught in traps baited with pistachio mummies compared with traps baited with almond mummies in almond orchards and pistachio orchards, respectively. Also, traps located in pistachio orchards caught 5.9 and 8.3 times more navel orangeworm than were trapped from almond orchards in 2009 and 2010, respectively. Implications for use of this novel baited trap in almond and pistachio orchard integrated pest management programs are discussed.     

Driving factors of the communities of phytophagous and predatory mites in a physic nut plantation and spontaneous plants associated

Seasonal changes in climate and plant diversity are known to affect the population dynamics of both pests and natural enemies within agroecosystems. In Brazil, spontaneous plants are usually tolerated in small-scale physic nut plantations over the year, which in turn may mediate interactions between pests and natural enemies within this agroecosystem. Here, we aimed to access the influence of seasonal variation of abiotic (temperature, relative humidity and rainfall) and biotic (diversity of spontaneous plants, overall richness and density of mites) factors on the communities of phytophagous and predatory mites found in a physic nut plantation and its associated spontaneous plants. Mite sampling was monthly conducted in dicotyledonous and monocotyledonous leaves of spontaneous plants as well as in physic nut shrubs over an entire year. In the dry season there was a higher abundance of phytophagous mites (Tenuipalpidae, Tarsonemidae and Tetranychidae) on spontaneous plants than on physic nut shrubs, while predatory mites (Phytoseiidae) showed the opposite pattern. The overall density of mites on spontaneous plants increased with relative humidity and diversity of spontaneous plants. Rainfall was the variable that most influenced the density of mites inhabiting physic nut shrubs. Agroecosystems comprising spontaneous plants associated with crops harbour a rich mite community including species of different trophic levels which potentially benefit natural pest control due to increased diversity and abundance of natural enemies.     

Comparison of pollinators and natural enemies: a meta‐analysis of landscape and local effects on abundance and richness in crops

To manage agroecosystems for multiple ecosystem services, we need to know whether the management of one service has positive, negative, or no effects on other services. We do not yet have data on the interactions between pollination and pest‐control services. However, we do have data on the distributions of pollinators and natural enemies in agroecosystems. Therefore, we compared these two groups of ecosystem service providers, to see if the management of farms and agricultural landscapes might have similar effects on the abundance and richness of both. In a meta‐analysis, we compared 46 studies that sampled bees, predatory beetles, parasitic wasps, and spiders in fields, orchards, or vineyards of food crops. These studies used the proximity or proportion of non‐crop or natural habitats in the landscapes surrounding these crops (a measure of landscape complexity), or the proximity or diversity of non‐crop plants in the margins of these crops (a measure of local complexity), to explain the abundance or richness of these beneficial arthropods. Compositional complexity at both landscape and local scales had positive effects on both pollinators and natural enemies, but different effects on different taxa. Effects on bees and spiders were significantly positive, but effects on parasitoids and predatory beetles (mostly Carabidae and Staphylinidae) were inconclusive. Landscape complexity had significantly stronger effects on bees than it did on predatory beetles and significantly stronger effects in non‐woody rather than in woody crops. Effects on richness were significantly stronger than effects on abundance, but possibly only for spiders. This abundance‐richness difference might be caused by differences between generalists and specialists, or between arthropods that depend on non‐crop habitats (ecotone species and dispersers) and those that do not (cultural species). We call this the ‘specialist‐generalist’ or ‘cultural difference’ mechanism. If complexity has stronger effects on richness than abundance, it might have stronger effects on the stability than the magnitude of these arthropod‐mediated ecosystem services. We conclude that some pollinators and natural enemies seem to have compatible responses to complexity, and it might be possible to manage agroecosystems for the benefit of both. However, too few studies have compared the two, and so we cannot yet conclude that there are no negative interactions between pollinators and natural enemies, and no trade‐offs between pollination and pest‐control services. Therefore, we suggest a framework for future research to bridge these gaps in our knowledge.     

Natural enemies of the maize cob borer,Mussidia nigrivenella (Lepidoptera: Pyralidae) in Benin,West Africa

Mussidia nigrivenella Ragonot is a pest of maize cobs in West Africa. It significantly reduces maize yields and grain quality, with quantitative losses of 2-25%at harvest, and up to 10-15% indirect losses due to an increase in storage pest infestation levels. Infestation by M. nigrivenella also significantly increased the susceptibility of maize to Aspergillus flavus infection and subsequent aflatoxin contamination. Surveys conducted in different agro-ecological zones of Benin on cultivated and wild host plants during 1994-1997 revealed one egg parasitoid, three larval parasitoids and one pupal parasitoid attacking M. nigrivenella. Egg parasitism was scarce on all host plants sampled and in all four agro-ecological zones. Parasitism by larval and pupal parasitoids was usually less than 10%, and varied with host plant species. Both larval and pupal parasitoids were rare or absent in cultivated maize fields. The solitary chalcidid pupal parasitoid, Antrocephalus crassipes Masi, was the predominant species, contributing approximately 53% of the observed mortality. Logistic regression analysis indicated that this parasitoid was more prevalent on fruits of Gardenia spp. (Rubiaceae) than on the other host plant species including maize used by M. nigrivenella, and was most abundant between February and September. The differences in parasitoid diversity and parasitism between Benin and other regions suggest that there are opportunities for biological control through introduction of exotic parasitoids or using the 'new association' approach, which uses natural enemies of closely related host species that occupy similar ecological niches to the target pest.