Parasitoid diversity reduces the variability in pest control services across time on farms

Recent declines in biodiversity have increased interest in the link between biodiversity and the provision and sustainability of ecosystem services across space and time. We mapped the complex network of interactions between herbivores and parasitoids to examine the relationship between parasitoid species richness, functional group diversity and the provision of natural pest control services. Quantitative food webs were constructed for 10 organic and 10 conventional farms. Parasitoid species richness varied from 26 to 58 species and we found a significant positive relationship between parasitoid species richness and temporal stability in parasitism rates. Higher species richness was associated with lower variation in parasitism rate. A functional group analysis showed significantly greater parasitoid species complementarity on organic farms, with on average more species in each functional group. We simulated parasitoid removal to predict whether organic farms experienced greater robustness of parasitism in the face of local extinctions. This analysis showed no consistent differences between the organic and conventional farm pairs in terms of loss of pest control service. Finally, it was found that the different habitats that make up each farm do not contribute equally to parasitoid species diversity, and that hedgerows produced more parasitoid species, significantly more so on organic farms.     

Functional benefits of predator species diversity depend on prey identity

Abstract.  1. Determining the functional significance of species diversity in natural enemy assemblages is a key step towards prediction of the likely impact of biodiversity loss on natural pest control processes. While the biological control literature contains examples in which increased natural enemy diversity hinders pest control, other studies have highlighted mechanisms where pest suppression is promoted by increased enemy diversity.
2. This study aimed to test whether increased predator species diversity results in higher rates of predation on two key, but contrasting, insect pest species commonly found in the rice ecosystems of south-east Asia.
3. Glasshouse experiments were undertaken in which four life stages of a planthopper ( Nilaparvata lugens ) and a moth ( Marasmia patnalis ) were caged with single or three-species combinations of generalist predators.
4. Generally, predation rates of the three-species assemblages exceeded expectation when attacking M. patnalis , but not when attacking N. lugens. In addition, a positive effect of increased predator species richness on overall predation rate was found with M. patnalis but not with N. lugens .
5. The results are consistent with theoretical predictions that morphological and behavioural differentiation among prey life stages promotes functional complementarity among predator species. This indicates that emergent species diversity effects in natural enemy assemblages are context dependent; they depend not only on the characteristics of the predators species, but on the identity of the species on which they prey.     

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

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Connecting scales: Achieving in‐field pest control from areawide and landscape ecology studies

Areawide management has a long history of achieving solutions that target pests, however, there has been little focus on the areawide management of arthropod natural enemies. Landscape ecology studies that show a positive relationship between natural enemy abundance and habitat diversity demonstrate landscape‐dependent pest suppression, but have not yet clearly linked their findings to pest management or to the suite of pests associated with crops that require control. Instead the focus has often been on model systems of single pest species and their natural enemies. We suggest that management actions to capture pest control from natural enemies may be forth coming if: (i) the suite of response and predictor variables focus on pest complexes and specific management actions; (ii) the contribution of “the landscape” is identified by assessing the timing and numbers of natural enemies immigrating and emigrating to and from the target crop, as well as pests; and (iii) pest control thresholds aligned with crop development stages are the benchmark to measure impact of natural enemies on pests, in turn allowing for comparison between study regions, and generalizations. To achieve pest control we will need to incorporate what has been learned from an ecological understanding of model pest and natural enemy systems and integrate areawide landscape management with in‐field pest management.     

A meta-analysis of crop pest and natural enemy response to landscape complexity

Many studies in recent years have investigated the relationship between landscape complexity and pests, natural enemies and/or pest control. However, no quantitative synthesis of this literature beyond simple vote-count methods yet exists. We conducted a meta-analysis of 46 landscape-level studies, and found that natural enemies have a strong positive response to landscape complexity. Generalist enemies show consistent positive responses to landscape complexity across all scales measured, while specialist enemies respond more strongly to landscape complexity at smaller scales. Generalist enemy response to natural habitat also tends to occur at larger spatial scales than for specialist enemies, suggesting that land management strategies to enhance natural pest control should differ depending on whether the dominant enemies are generalists or specialists. The positive response of natural enemies does not necessarily translate into pest control, since pest abundances show no significant response to landscape complexity. Very few landscape-scale studies have estimated enemy impact on pest populations, however, limiting our understanding of the effects of landscape on pest control. We suggest focusing future research efforts on measuring population dynamics rather than static counts to better characterise the relationship between landscape complexity and pest control services from natural enemies.     

Organic farming expansion drives natural enemy abundance but not diversity in vineyard‐dominated landscapes

Organic farming is seen as a prototype of ecological intensification potentially able to conciliate crop productivity and biodiversity conservation in agricultural landscapes. However, how natural enemies, an important functional group supporting pest control services, respond to organic farming at different scales and in different landscape contexts remain unclear. Using a hierarchical design within a vineyard‐dominated region located in southwestern France, we examine the independent effects of organic farming and semi‐natural habitats at the local and landscape scales on natural enemies. We show that the proportion of organic farming is a stronger driver of species abundance than the proportion of semi‐natural habitats and is an important facet of landscape heterogeneity shaping natural enemy assemblages. Although our study highlights a strong taxonomic group‐dependency about the effect of organic farming, organic farming benefits to dominant species while rare species occur at the same frequency in the two farming systems. Independently of farming systems, enhancing field age, reducing crop productivity, soil tillage intensity, and pesticide use are key management options to increase natural enemy biodiversity. Our study indicates that policies promoting the expansion of organic farming will benefit more to ecological intensification strategies seeking to enhance ecosystem services than to biodiversity conservation.     

农田景观格局对害虫天敌生态控害功能的调控作用

如何提升农田景观的生态服务功能是当前景观生态学和害虫防治学的前沿研究热点.基于区域农田景观格局可显著调节植物-植食性昆虫-天敌之间相互关系的理论基础,本文系统概括总结了农田景观格局及其变化对农田重要天敌多样性与生态控害功能的影响.从不同天敌类群的角度,分析了农田景观格局对捕食性天敌、寄生性天敌、有益微生物及其生态控害功能的调控作用.同时,对优化农田景观作物布局、采取合理的植被生境管理策略进行了展望.本文可为应用植被生境管理提升农田景观中天敌的生态系统服务功能提供参考,进一步促进区域生态农业响应"化学农药减量施用"的战略目标提供理论支撑.     

Interaction effects between local flower richness and distance to natural woodland on pest and beneficial insects in apple orchards

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Forest bolsters bird abundance,pest control and coffee yield

Efforts to maximise crop yields are fuelling agricultural intensification, exacerbating the biodiversity crisis. Low‐intensity agricultural practices, however, may not sacrifice yields if they support biodiversity‐driven ecosystem services. We quantified the value native predators provide to farmers by consuming coffee's most damaging insect pest, the coffee berry borer beetle (Hypothenemus hampei). Our experiments in Costa Rica showed birds reduced infestation by ~ 50%, bats played a marginal role, and farmland forest cover increased pest removal. We identified borer‐consuming bird species by assaying faeces for borer DNA and found higher borer‐predator abundances on more forested plantations. Our coarse estimate is that forest patches doubled pest control over 230 km² by providing habitat for ~ 55 000 borer‐consuming birds. These pest‐control services prevented US$75–US$310 ha‐year⁻¹ in damage, a benefit per plantation on par with the average annual income of a Costa Rican citizen. Retaining forest and accounting for pest control demonstrates a win–win for biodiversity and coffee farmers.     

Response diversity determines the resilience of ecosystems to environmental change

A growing body of evidence highlights the importance of biodiversity for ecosystem stability and the maintenance of optimal ecosystem functionality. Conservation measures are thus essential to safeguard the ecosystem services that biodiversity provides and human society needs. Current anthropogenic threats may lead to detrimental (and perhaps irreversible) ecosystem degradation, providing strong motivation to evaluate the response of ecological communities to various anthropogenic pressures. In particular, ecosystem functions that sustain key ecosystem services should be identified and prioritized for conservation action. Traditional diversity measures (e.g. ‘species richness’) may not adequately capture the aspects of biodiversity most relevant to ecosystem stability and functionality, but several new concepts may be more appropriate. These include ‘response diversity’, describing the variation of responses to environmental change among species of a particular community. Response diversity may also be a key determinant of ecosystem resilience in the face of anthropogenic pressures and environmental uncertainty. However, current understanding of response diversity is poor, and we see an urgent need to disentangle the conceptual strands that pervade studies of the relationship between biodiversity and ecosystem functioning. Our review clarifies the links between response diversity and the maintenance of ecosystem functionality by focusing on the insurance hypothesis of biodiversity and the concept of functional redundancy. We provide a conceptual model to describe how loss of response diversity may cause ecosystem degradation through decreased ecosystem resilience. We explicitly explain how response diversity contributes to functional compensation and to spatio‐temporal complementarity among species, leading to long‐term maintenance of ecosystem multifunctionality. Recent quantitative studies suggest that traditional diversity measures may often be uncoupled from measures (such as response diversity) that may be more effective proxies for ecosystem stability and resilience. Certain conclusions and recommendations of earlier studies using these traditional measures as indicators of ecosystem resilience thus may be suspect. We believe that functional ecology perspectives incorporating the effects and responses of diversity are essential for development of management strategies to safeguard (and restore) optimal ecosystem functionality (especially multifunctionality). Our review highlights these issues and we envision our work generating debate around the relationship between biodiversity and ecosystem functionality, and leading to improved conservation priorities and biodiversity management practices that maximize ecosystem resilience in the face of uncertain environmental change.     

Biocontrol potential varies with changes in biofuel–crop plant communities and landscape perenniality

We examined the potential local‐ and landscape‐level impacts of different biofuel production systems on biocontrol, an important service provided by arthropod natural enemies. Specifically, we sampled natural enemies with sweep nets and measured predation of sentinel pest eggs in stands of corn, switchgrass and mixed prairie in Michigan and Wisconsin (total n=40 for natural enemy sampling, n=60 for egg predation), relating them to crop type, forb cover and diversity, and the composition and heterogeneity of the surrounding landscape. Grasslands with intermediate levels of forb cover and flower diversity supported two‐orders of magnitude more natural enemy biomass, fourfold more natural enemy families, and threefold greater rates of egg predation than corn. Data suggest this was in part due to a general increase in biomass, richness and predation in perennial grasslands compared with corn, combined with a positive effect of intermediate levels of forb cover and flower diversity. Specifically, natural enemy biomass and family richness showed hump‐shaped relationships to forb cover that peaked in sites with 5–25% forbs, while egg predation increased with floral diversity. At the landscape scale, both natural‐enemy biomass and egg predation increased with the area of forest in the landscape, and egg predation almost doubled as the area of herbaceous, perennial habitats within 1.5 km of study sites increased. Our results suggest that floristically diverse, perennial grasslands support diverse and abundant predator communities that contribute to natural pest suppression. In addition, large‐scale production of biofuel crops could positively or negatively affect biocontrol services in agricultural landscapes through associated changes in the area of perennial habitats. Biofuel landscapes that incorporate perennial grasslands could support a variety of beneficial organisms and ecosystem services in addition to producing biomass.     

Effects of biodiversity in agricultural landscapes on the protective microbiome of insects – a review

Symbiotic bacteria in herbivorous insects can have strong beneficial impacts on their host's survival, including conferring resistance to natural enemies such as parasitoid wasps or pathogens, while also imposing energetic costs on the host, resulting in cost‐benefit trade‐offs. Whether these trade‐offs favour the hosting of symbionts depends on the growth environment of the herbivore. Long‐term experimental grassland studies have shown that increasing plant species richness leads to an increased diversity of associated herbivores and their natural enemies. Such a change in natural enemy diversity, related to changes in plant diversity, could also drive changes in the community of symbionts hosted by the herbivorous insects. Aphids are one model system for studying symbionts in insects, and effects of host‐plant species and diversity on aphid‐symbiont interactions have been documented. Yet, we still understand little of the mechanisms underlying such effects. We review the current state of knowledge of how biodiversity can impact aphid‐symbiont communities and the underlying drivers. Then, we discuss this in the framework of sustainable agriculture, where increased plant biodiversity, in the form of wildflower strips, is used to recruit natural enemies to crop fields for their pest control services. Although aphid symbionts have the potential to reduce biological control effectiveness through conferring protection for the host insect, we discuss how increasing plant and natural enemy biodiversity can mitigate these effects and identify future research opportunities. Understanding how to promote beneficial interactions in ecological systems can help in the development of more sustainable agricultural management strategies.     

农业景观生物多样性与害虫生态控制

现代农业的一个重要特征就是人类对农田生态系统的干扰强度及频率不断增加,严重影响农业景观的结构及其生物多样性.农业景观结构的变化及其生物多样性的丧失,必然引起生态系统服务功能的弱化,不利于实施以保护自然天敌为主的害虫生态控制.农业的集约化经营导致自然生境破碎化,减少了农业景观的复杂性,使得作物和非作物变成一种相对离散化的生境类型和镶嵌的景观格局;破碎化的生境不仅会减少某些物种的丰度,还会影响物种之间的相互关系及生物群落的多样性和稳定性.非作物生境类型如林地、灌木篱墙、田块边缘区、休耕地和草地等,是一种比较稳定的异质化环境.非作物生境较少受到干扰,可以为寄生性和捕食性节肢动物提供适宜的越冬或避难场所以及替代猎物、花粉和花蜜等资源,因此,非作物生境有利于自然天敌的栖息和繁衍,也有利于它们迁入邻近的作物生境中对害虫起到调节和控制作用.景观的格局-过程-尺度影响农田生物群落物种丰富度、多度、多样性以及害虫与天敌之间的相互作用.从区域农业景观系统的角度出发,运用景观生态学的理论和方法来研究作物、害虫、天敌等组分在不同斑块之间的转移过程和变化规律,揭示害虫在较大尺度和具有异质性的空间范围内的灾变机理,可为利用农业景观生物多样性来保护农田自然天敌,实施害虫的区域性生态控制提供新的研究思路和手段.     

Relationships between biodiversity and biological control in agroecosystems: Current status and future challenges

Agricultural systems around the world are faced with the challenge of providing for the demands of a growing human population. To meet this demand, agricultural systems have intensified to produce more crops per unit area at the expense of greater inputs. Agricultural intensification, while yielding more crops, generally has detrimental impacts on biodiversity. However, intensified agricultural systems often have fewer pests than more “environmentally-friendly” systems, which is believed to be primarily due to extensive pesticide use on intensive farms. In turn, to be competitive, less-intensive agricultural systems must rely on biological control of pests. Biological pest control is a complex ecosystem service that is generally positively associated with biodiversity of natural enemy guilds. Yet, we still have a limited understanding of the relationships between biodiversity and biological control in agroecosystems, and the mechanisms underlying these relationships. Here, we review the effects of agricultural intensification on the diversity of natural enemy communities attacking arthropod pests and weeds. We next discuss how biodiversity of these communities impacts pest control, and the mechanisms underlying these effects. We focus in particular on novel conceptual issues such as relationships between richness, evenness, abundance, and pest control. Moreover, we discuss novel experimental approaches that can be used to explore the relationships between biodiversity and biological control in agroecosystems. In particular, we highlight new experimental frontiers regarding evenness, realistic manipulations of biodiversity, and functional and genetic diversity. Management shifts that aim to conserve diversity while suppressing both insect and weed pests will help growers to face future challenges. Moreover, a greater understanding of the interactions between diversity components, and the mechanisms underlying biodiversity effects, would improve efforts to strengthen biological control in agroecosystems.     

Natural disturbance impacts on ecosystem services and biodiversity in temperate and boreal forests

In many parts of the world forest disturbance regimes have intensified recently, and future climatic changes are expected to amplify this development further in the coming decades. These changes are increasingly challenging the main objectives of forest ecosystem management, which are to provide ecosystem services sustainably to society and maintain the biological diversity of forests. Yet a comprehensive understanding of how disturbances affect these primary goals of ecosystem management is still lacking. We conducted a global literature review on the impact of three of the most important disturbance agents (fire, wind, and bark beetles) on 13 different ecosystem services and three indicators of biodiversity in forests of the boreal, cool‐ and warm‐temperate biomes. Our objectives were to (i) synthesize the effect of natural disturbances on a wide range of possible objectives of forest management, and (ii) investigate standardized effect sizes of disturbance for selected indicators via a quantitative meta‐analysis. We screened a total of 1958 disturbance studies published between 1981 and 2013, and reviewed 478 in detail. We first investigated the overall effect of disturbances on individual ecosystem services and indicators of biodiversity by means of independence tests, and subsequently examined the effect size of disturbances on indicators of carbon storage and biodiversity by means of regression analysis. Additionally, we investigated the effect of commonly used approaches of disturbance management, i.e. salvage logging and prescribed burning. We found that disturbance impacts on ecosystem services are generally negative, an effect that was supported for all categories of ecosystem services, i.e. supporting, provisioning, regulating, and cultural services (P < 0.001). Indicators of biodiversity, i.e. species richness, habitat quality and diversity indices, on the other hand were found to be influenced positively by disturbance (P < 0.001). Our analyses thus reveal a ‘disturbance paradox’, documenting that disturbances can put ecosystem services at risk while simultaneously facilitating biodiversity. A detailed investigation of disturbance effect sizes on carbon storage and biodiversity further underlined these divergent effects of disturbance. While a disturbance event on average causes a decrease in total ecosystem carbon by 38.5% (standardized coefficient for stand‐replacing disturbance), it on average increases overall species richness by 35.6%. Disturbance‐management approaches such as salvage logging and prescribed burning were neither found significantly to mitigate negative effects on ecosystem services nor to enhance positive effects on biodiversity, and thus were not found to alleviate the disturbance paradox. Considering that climate change is expected to intensify natural disturbance regimes, our results indicate that biodiversity will generally benefit from such changes while a sustainable provisioning of ecosystem services might come increasingly under pressure. This underlines that disturbance risk and resilience require increased attention in ecosystem management in the future, and that new approaches to addressing the disturbance paradox in management are needed.     

Multitrophic diversity effects of network degradation

Predicting the functional consequences of biodiversity loss in realistic, multitrophic communities remains a challenge. No existing biodiversity–ecosystem function study to date has simultaneously incorporated information on species traits, network topology, and extinction across multiple trophic levels, while all three factors are independently understood as critical drivers of post‐extinction network structure and function. We fill this gap by comparing the functional consequences of simulated species loss both within (monotrophic) and across (bitrophic) trophic levels, in an ecological interaction network estimated from spatially explicit field data on tropical fecal detritus producer and consumers (mammals and dung beetles). We simulated trait‐ordered beetle and mammal extinction separately (monotrophic extinction) and the coextinction of beetles following mammal loss (bitrophic extinction), according to network structure. We also compared the diversity effects of bitrophic extinction models using a standard monotrophic function (the daily production or consumption of fecal detritus) and a unique bitrophic functional metric (the proportion of daily detritus production that is consumed). We found similar mono‐ and bitrophic diversity effects, regardless of which species traits were used to drive extinctions, yet divergent predictions when different measures of function were used. The inclusion of information on network structure had little apparent effect on the qualitative relationship between diversity and function. These results contribute to our growing understanding of the functional consequences of biodiversity from real systems and underscore the importance of species traits and realistic functional metrics to assessments of the ecosystem impacts of network degradation through species loss.     

Interactions between human behaviour and ecological systems

Research on the interactions between human behaviour and ecological systems tends to focus on the direct effects of human activities on ecosystems, such as biodiversity loss. There is also increasing research effort directed towards ecosystem services. However, interventions to control people's use of the environment alter the incentives that natural resource users face, and therefore their decisions about resource use. The indirect effects of conservation interventions on biodiversity, modulated through human decision-making, are poorly studied but are likely to be significant and potentially counterintuitive. This is particularly so where people are dependent on multiple natural resources for their livelihoods, when both poverty and biodiversity loss are acute. An inter-disciplinary approach is required to quantify these interactions, with an understanding of human decision-making at its core; otherwise, predictions about the impacts of conservation policies may be highly misleading.     

Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition

Global change, especially land‐use intensification, affects human well‐being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real‐world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land‐use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land‐use objectives. We found that indirect land‐use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land‐use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land‐use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast‐growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.     

Sustainable pest regulation in agricultural landscapes: a review on landscape composition, biodiversity and natural pest control

Agricultural intensification has resulted in a simplification of agricultural landscapes by the expansion of agricultural land, enlargement of field size and removal of non-crop habitat. These changes are considered to be an important cause of the rapid decline in farmland biodiversity, with the remaining biodiversity concentrated in field edges and non-crop habitats. The simplification of landscape composition and the decline of biodiversity may affect the functioning of natural pest control because non-crop habitats provide requisites for a broad spectrum of natural enemies, and the exchange of natural enemies between crop and non-crop habitats is likely to be diminished in landscapes dominated by arable cropland. In this review, we test the hypothesis that natural pest control is enhanced in complex patchy landscapes with a high proportion of non-crop habitats as compared to simple large-scale landscapes with little associated non-crop habitat. In 74% and 45% of the studies reviewed, respectively, natural enemy populations were higher and pest pressure lower in complex landscapes versus simple landscapes. Landscape-driven pest suppression may result in lower crop injury, although this has rarely been documented. Enhanced natural enemy activity was associated with herbaceous habitats in 80% of the cases (e.g. fallows, field margins), and somewhat less often with wooded habitats (71%) and landscape patchiness (70%). The similar contributions of these landscape factors suggest that all are equally important in enhancing natural enemy populations. We conclude that diversified landscapes hold most potential for the conservation of biodiversity and sustaining the pest control function.     

Effects of crop species richness on pest-natural enemy systems based on an experimental model system using a microlandscape

The relationship between crop richness and predator-prey interactions as they relate to pest-natural enemy systems is a very important topic in ecology and greatly affects biological control services. The effects of crop arrangement on predator-prey interactions have received much attention as the basis for pest population management. To explore the internal mechanisms and factors driving the relationship between crop richness and pest population management, we designed an experimental model system of a microlandscape that included 50 plots and five treatments. Each treatment had 10 repetitions in each year from 2007 to 2010. The results showed that the biomass of pests and their natural enemies increased with increasing crop biomass and decreased with decreasing crop biomass; however, the effects of plant biomass on the pest and natural enemy biomass were not significant. The relationship between adjacent trophic levels was significant (such as pests and their natural enemies or crops and pests), whereas non-adjacent trophic levels (crops and natural enemies) did not significantly interact with each other. The ratio of natural enemy/pest biomass was the highest in the areas of four crop species that had the best biological control service. Having either low or high crop species richness did not enhance the pest population management service and lead to loss of biological control. Although the resource concentration hypothesis was not well supported by our results, high crop species richness could suppress the pest population, indicating that crop species richness could enhance biological control services. These results could be applied in habitat management aimed at biological control, provide the theoretical basis for agricultural landscape design, and also suggest new methods for integrated pest management.