基于服务功能的昆虫生态调控理论

鉴于昆虫在植物传粉授精、害虫生物控制、土壤有机物分解中提供多种生态系统服务功能,本文在害虫生态调控、区域性害虫生态调控与生境管理的基础上,进一步提出基于多种生态服务功能的农田景观昆虫生态调控理论、方法与实践。认为:昆虫管理不仅仅是害虫的管理,还应包括有益昆虫(如传粉昆虫、天敌昆虫、分解昆虫)的管理,这种管理应从单一农田生态系统扩展到农田景观生态系统,充分考虑农田景观中昆虫的传粉功能、生物控害功能和分解功能,通过对功能植物、作物与非作物生境的空间布局以及时间序列上的生态设计,从空间上明确昆虫(包括害虫、天敌、传粉昆虫、分解昆虫)在不同生境中的转移扩散动态,从时间上掌握昆虫在不同寄主植物与非作物生境上的演替过程,从技术上着重发挥有利于昆虫的传粉功能、生物控害功能和分解功能的综合措施,在研究方法上突出使用稳定同位素、生态能量学、化学生态学等定量分析手段,研究景观区域内中"植物-昆虫"互作过程及其生态调控措施的作用,寻求不同时空条件下控害保益的关键措施,设计和组装出维持多功能的农田景观昆虫生态调控技术体系,创造有利于天敌控害、蜜蜂传粉、土壤分解的环境条件,以发挥昆虫类群在农田景观中最大的生态服务功能。     

生境管理——保护性生物防治的发展方向

生境管理是近年来保护性生物防治的重要研究方向,也是利用农业景观格局进行生物防治的重要策略。生境管理是指从农田景观的角度,在大时空尺度范围内进行多种作物与非作物生境的设计与布局,创造有利于天敌的环境条件,抑制害虫种群发生,达到减小环境污染、增强农业生态系统的控害保益功能,最终实现害虫种群控制的可持续性。景观尺度下的生境管理不仅强调单一农田生物控害作用,而是以多生境农业景观整体布局为指导,探索各种生境功能的整合利用,以发挥各种生境最大的生物控害潜能,为实现多目标生态服务价值管理提供重要的理论基础和现实依据。本文系统地总结了生境管理的研究内容与实现途径,论述了农业景观格局与过程对害虫种群控制的机理,并对作为保护性生物防治发展方向的生境管理研究趋势进行了展望。     

多尺度空间下害虫生态调控理论与应用

不同尺度空间下农田生态系统具有不同的生境斑块组成结构,尺度性也是生态系统的重要特征之一.近年来,北美和欧洲等地区用农田生境管理与区域性景观设计相结合的研究方法实施多尺度空间下害虫生态调控,实现复合生态系统服务.其核心思想是以大区域景观设计和农田作物布局与农事管理的有机结合,通过农业景观格局的空间配置,调节种植模式、管理技术、乃至改变农业景观格局的空间配置等以切断害虫种群的生活史,建立和恢复天敌种群库与转移通道,从而最大程度地提高农业生态系统自身的控害功能.近年来,北美和欧洲对多尺度空间下农业复合生态系统服务功能都做了大量工作,尤其是田间尺度与景观尺度相结合的研究方法更是当前生境管理研究的重要内容.本文总结了多尺度空间下生态系统环境条件与天敌种群间的作用机制及假说,包括田间尺度上主要通过轮作与间套作、覆盖作物、减免耕及发展有机农业等方式提高天敌种群,景观尺度上通过生境斑块的空间配置来改变植物资源布局,最终提高天敌的控害作用.以期为深入地解析景观格局及复杂性对生物多样性的影响,揭示农业景观变化对昆虫种间关系的作用机制,在实践上为利用农田景观格局控制害虫种群发生提供新的途径与方法.     

半自然农田边界与相邻农田步甲和蜘蛛的时空分布

分别于小麦、玉米收获前后,采用陷阱法调查了华北地区典型农业景观中具有不同植被结构的农田边界及其相邻农田中两类重要天敌类群步甲和蜘蛛的多样性.通过比较农田生境及相邻农田边界间两类天敌群落的时空分布格局及其与相邻半自然生境植被群落的相关关系,探讨半自然农田边界对两类天敌类群的保护作用.结果表明: 整个取样季节农田边界处蜘蛛的多度显著高于农田内部;而步甲多样性在农田与边界间无显著性差异,仅呈现不同的群落结构;作物收获后蜘蛛分科数在边界处的增加以及在农田的减少,显示了蜘蛛在农田和边界之间的迁移活动.边界植被结构对蜘蛛和步甲多样性有不同影响:边界较高的草本层盖度和较低的乔木层盖度有利于增加农田中某些步甲优势种的多样性;而较高的草本层盖度有利于增加皿蛛科蜘蛛的多样性.因此,半自然生境的存在可以通过天敌在农田和边界之间的迁移运动促进农田天敌多样性的维持;但不同类型半自然生境植被群落结构可能影响其对不同天敌群落多样性的维持和保护作用.为促进农业景观对天敌的保护作用,提高其害虫控制功能,需要深入了解不同天敌的生境需求及食物需求,精心设计有利于天敌多样性维持的半自然生境.     

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.     

区域性农田景观格局对麦蚜及其天敌种群的生态学效应

明确农田景观格局对害虫及其天敌种群的生态学效应,是开展区域性害虫生态调控的基础.以前的研究大多集中于小空间尺度下、单个景观因子对昆虫种群的作用,而从省级范围的大空间尺度、多个景观因子的分析很少.本文以山东省区域性小麦种植区为研究对象,基于遥感影像和土地覆盖分类数据以及田间调查的昆虫种群数据,分析了景观组成类型(component type)、构成比例(component proportions)和形状结构(shape structure)多因子对麦蚜及其天敌寄生蜂和瓢虫种群的综合作用.结果发现,农田景观组成类型中斑块类型(patch type)越多,越利于麦蚜和天敌瓢虫种群数量的增长;且斑块密度(patch density)越大,越利于麦蚜寄生蜂和天敌瓢虫数量的增加;景观形状结构中边界密度(edge density)越高,也越利于麦蚜寄生蜂和天敌瓢虫种群数量的增加.进一步定量评估了农田景观组成类型、构成比例和形状结构对麦蚜及其天敌种群影响的作用大小.结果表明,三类景观格局因子对麦蚜影响较小,权重为9.81%;而对麦蚜寄生蜂的影响权重为25.87%;对天敌瓢虫种群高达47.86%.本研究清楚地表明,通过优化农田景观中作物与非作物生境布局,可直接调节和增加天敌昆虫种类与数量,有效控制和减少小麦蚜虫的种群数量,从而提高区域性农田景观中天敌昆虫的生物控害服务功能.     

功能植物苣荬菜的特征及其应用潜能

苣荬菜Sonchus arvensis L.在我国广泛分布,具有耐干旱、耐盐碱、适应性强、管理简便、花期长的特点。我们于2018年9月最先在山东发现苣荬菜上涵养着大量的天敌昆虫和传粉昆虫。经过进一步调研,显示苣荬菜与周围作物(玉米、小麦)无共同害虫,其花期7-10月,与玉米生长同期,可为玉米田天敌昆虫提供持续的营养补充,维持天敌昆虫种群;同时也是一种优质的蜜粉源植物,符合功能植物的基本特征。本文综述了苣荬菜作为华北玉米田功能植物的特征,建议通过在玉米田边通过种植苣荬菜可作为玉米害虫生态控害的手段,以实现维持天敌生物控害、减少化学农药投入、增加收入、改善农田生态环境和美丽乡村等目标。     

农田景观格局变化对昆虫的生态学效应

景观格局变化是全球变化的一个重要内容。农田是由人类赖以生存所种植的人工栽培作物组成的生态系统。在该景观系统中,多种植物-害虫-天敌相互作用、相互制约,形成有机整体。研究农田景观格局对害虫和天敌种群动态影响,不仅在害虫生物防治的实践中有重要意义,而且对于揭示人类活动对生物多样性结构与功能的影响,阐明农田景观中生物多样性整合、维持机理有重大的理论意义。本文从农田景观格局的"质、量、形、度"4个方面,系统地论述了农田景观格局变化对害虫和天敌的作用,分析了农田景观的生物控制服务功能,指出了未来研究发展的方向。     

区域农田景观格局对麦田天敌瓢虫群落的影响

【目的】明确农田景观格局对麦田天敌瓢虫种群的影响,为开展区域性害虫生态调控提供理论依据。【方法】以山东省22个县市区域的小麦种植区为研究对象,基于遥感影像与土地覆盖分类数据以及田间调查的瓢虫种群数据,计算景观格局指数,使用负二项分布的广义线性模型从农田景观、非作物生境景观和区域景观3个方面分析区域农田景观格局对麦田天敌瓢虫群落的影响。【结果】麦田瓢虫种群数量与草地的平均斑块面积(mean patch area,AREA_MN)和面积加权平均斑块分维数(area-weighted mean patch fractal dimension,FRAC_AM)、区域景观的斑块丰富度密度(patch richness density,PRD)呈正相关,与非作物生境的面积加权平均几何最邻近距离(areaweighted mean Euclidean nearest neighbor distance,ENN_AM)呈负相关。草地、聚集的非作物生境以及多样性的区域景观有利于天敌瓢虫种群数量的增加。使用草地的平均斑块面积和非作物生境的面积加权平均几何最邻近距离可以预测瓢虫的发生量。【结论】作为非作物生境的草地、非作物生境的空间分布及区域景观的多样性是影响麦田天敌瓢虫发生的重要因素。     

定量评价天敌昆虫控害功能的稳定同位素方法

天敌在区域性多作物农田景观系统中辗转捕食害虫,对调节和控制害虫种群数量发挥着非常重要的作用。明确天敌昆虫的食物来源和扩散规律是定量评价其控害功能的重要环节。其中,稳定同位素标记方法是追溯生物个体的食物来源和探究其运动规律的重要手段。本文首先介绍该标记方法的基本原理,接着以天敌昆虫龟纹瓢虫Propylea japonica(Thunberg)为例,应用稳定同位素碳标记方法追溯其在棉花和玉米农田景观系统中取食来源与食物比例;最后叙述了稳定同位素方法的应用前景和存在的问题。     

农田非作物生境调控与害虫综合治理

就害虫综合治理的研究而言,农田生态系统可以区分出作物生境和邻近作物的非作物生境。昆虫从作物生境迁移到非作物生境,与作物生境缺乏食物或受人类栽培活动干扰而引起的自然迁移、转换寄主和寻求庇护场所有关。许多研究表明,与特定作物田块相联系的植被类型和结构可影响害虫及其天敌迁居的种类、数量和时间。因此,我们可以通过改变大田周围非作物生境的植被组成及特征来调控农业生态系统中害虫与天敌的相互关系,提高天敌对害虫的控制效能。由于景观的空间格局对节肢动物的生物学特性有直接或间接的影响,所以,以景观为单元要比以同类作物的田块为单元更加适于害虫综合治理的研究和实施。在现代农业景观区域内重新引入和相嵌一些非栽培植物或廊道,可为众多有益节肢动物的繁殖、取食和避害提供多种类型的非作物栖境。     

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.     

Mechanisms for flowering plants to benefit arthropod natural enemies of insect pests： Prospects for enhanced use in agriculture

Reduction of noncrop habitats, intensive use of pesticides and high levels of disturbance associated with intensive crop production simplify the farming landscape and bring about a sharp decline of biodiversity. This, in turn, weakens the biological control ecosystem service provided by arthropod natural enemies. Strategic use of flowering plants to enhance plant biodiversity in a well-targeted manner can provide natural enemies with food sources and shelter to improve biological control and reduce dependence on chemical pesticides. This article reviews the nutritional value of various types of plant-derived food for natural enemies, possible adverse effects on pest management, and the practical application of flowering plants in orchards, vegetables and field crops, agricultural systems where most research has taken place. Prospects for more effective use of flowering plants to maximize biological control of insect pests in agroecosystem are good but depend up on selection of optimal plant species based on information on the ecological mechanisms by which natural enemies are selectively favored over pest species.     

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.     

Prospects for improved off-crop habitat management for pollen beetle control in oilseed rape

There is an urgent need to develop sustainable and environmentally benign integrated pest management (IPM) strategies for arable crops. The enhancement and manipulation of naturally occurring populations of the natural enemies of crop pests through habitat management for ‘conservation biological control’, as well as habitat management to manipulate populations of the pests themselves, have the potential to become major components of successful IPM strategies. We review the studies that have contributed to our current understanding of how the crop margin, local landscape, and regional landscape can influence pollen beetle Brassicogethes aeneus (syn. Meligethes aeneus) (Coleoptera: Nitidulidae) abundance and damage to oilseed rape crops (Brassica napus), and the efficacy of their natural enemies. We also discuss how habitat management across these multiple scales may improve pollen beetle control, reducing the need for insecticide use and contributing towards sustainable production of this important crop which is grown on increasing areas for both food and fuel.     

Landscape-scale forest cover increases the abundance of Drosophila suzukii and parasitoid wasps

Agricultural landscapes rich in natural and semi-natural habitats promote biodiversity and important ecosystem services for crops such as pest control. However, semi-natural habitats may fail to deliver these services if agricultural pests are disconnected from the available pool of natural enemies, as may be the case with invasive species. This study aimed to provide insights into the relationship between landscape complexity and the abundance of the recently established invasive pest species Drosophila suzukii and a group of natural enemies (parasitoid wasps), which contain species that parasitize D. suzukii in native and invaded ecosystems. The importance of landscape complexity was examined at two spatial scales. At the field scale, the response to introduction of wildflower strips was analysed, while the relationship with forest cover was assessed at the landscape scale. Half of the surveys were done next to blueberry crops (Vaccinium corymbosum), the other half was done in landscapes without fruit crops to examine effects of D. suzukii host presence. As expected, the number of observed parasitoid wasps increased with amount of forest surrounding the blueberry fields, but the number of D. suzukii individuals likewise increased with forest cover. Establishment of wildflower strips did not significantly affect the abundance of D. suzukii or parasitoid wasps and insect phenology was similar in landscapes with and without blueberry crops. This suggests that D. suzukii is enhanced by landscape complexity and is largely unlinked from the species group that, in its native range, hosts key natural enemies. Although management practices that rely on enhancing natural enemies through habitat manipulations can contribute to the long-term stability of agroecosystems and to control agricultural pests, other control measures may still be necessary in the short term to counteract the benefits obtained by D. suzukii from natural habitats.     

Habitat manipulation to mitigate the impacts of invasive arthropod pests

Exotic invaders are some of the most serious insect pests of agricultural crops around the globe. Increasingly, the structure of landscape and habitat is recognized as having a major influence on both insect pests and their natural enemies. Habitat manipulation that aims at conserving natural enemies can potentially contribute to safer and more effective control of invasive pests. In this paper, we review habitat management experiments, published during the last 10 years, which have aimed to improve biological control of invasive pests. We then discuss during what conditions habitat management to conserve natural enemies is likely to be effective and how the likelihood of success of such methods can be improved. We finally suggest an ecologically driven research agenda for habitat management programmes.