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。     

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.     

麦蚜和寄生蜂对农业景观格局的响应及其关键景观因子分析

大量研究表明多样性的农业景观格局能够影响蚜虫及寄生蜂的分布。本文利用并设计了两种尺度的麦田农业景观格局,依据麦蚜种群发生特点,分为迁入期、增长期与高峰期三个时期,论述了不同尺度下农业景观元素对麦蚜及寄生蜂系统的影响,通过逐步回归筛选了不同时期麦蚜及寄生蜂分布的关键景观元素,最后通过CANOCO软件模拟了麦蚜及寄生蜂的分布排序格局。结果表明不同景观因子对麦蚜及寄生蜂种群影响不同,迁入期两种尺度下裸地最有利于两种有翅蚜的迁飞入田,塑料大棚对有翅蚜种群的入田有抑制作用。增长期草地与林地生境对麦蚜种群增长率促进最大,塑料大棚同样抑制了麦蚜的种群增长率;但塑料大棚、草地与林地对蚜茧蜂的种群增长率有促进作用,非麦类作物生境对蚜茧蜂种群增长率抑制作用最大;草地与林地有利于重寄生蜂的种群增长率;高峰期裸地比例大的农业景观下麦蚜的最大种群密度较大,草地与林地对蚜茧蜂与重寄生蜂的最大种群密度均有促进作用。两种尺度下的研究结果一致。不同麦物种的对不同景观元素反应与形态学与生活史特征有关,而且景观结构中特定的植物种类、非作物植物的密度与物候期都可能影响寄生蜂群落的多样性与功能。     

区域性景观组成对玉米苗期捕食性天敌的生态学效应

【目的】明确区域性景观组成对玉米苗期捕食性天敌的生态学效应,可为开展玉米田有害生物生态调控提供理论基础。【方法】以山东省92块玉米田为研究区域,基于玉米苗期捕食性天敌种群数量、试验点的遥感影像和土地覆盖类型分类数据,运用混合效应模型分析了耕地、居住工业交通、草地、林地、水体5种土地覆盖类型的面积比例对玉米苗期捕食性天敌种群数量的影响。【结果】共调查到3 744头捕食性天敌,其中草间小黑蛛Erigonidium graminicolum(55.29%),龟纹瓢虫Propylaea japonica(25.32%),小花蝽Orius similis(6.73%),八斑球腹蛛Theridion octomaculatum(4.01%),异色瓢虫Harmonia axyridis(3.58%)和三突花蛛Misumenops tricuspidatus(3.47%)为主要捕食性天敌。发现林地和水体生境对龟纹瓢虫的种群数量有显著的正效应,草地和水体生境对三突花蛛的种群数量有显著的正效应,水体和林地生境对捕食性天敌的总数量有显著正效应。【结论】华北农田景观中非作物生境作为捕食性天敌的资源库,在玉米苗期捕食性天敌维持中起重要作用。     

麦蚜与龟纹瓢虫在麦田的空间格局分析

【目的】为保护麦田天敌并合理利用天敌对麦蚜进行控制。【方法】通过田间网格式取样调查,应用生态位和地统计学方法研究了麦田中麦蚜及其主要天敌龟纹瓢虫Proplaea japonica的时空分布格局。【结果】麦蚜及瓢虫时间序列上的种群动态分析显示,麦田中龟纹瓢虫发生期比麦蚜滞后,麦蚜与龟纹瓢虫的种群数量动态趋势相似,种群相关系数为0.930,表明龟纹瓢虫与麦蚜种群呈明显的跟随关系。在空间上,麦蚜和龟纹瓢虫两者的生态位重叠度随着麦蚜的发生量增多而增高,最高达0.52。麦蚜与龟纹瓢虫的空间格局在时间序列上,随着种群数量改变而变化。在低种群密度下,麦蚜的半变异函数为线性模型,呈随机分布;在高密度下,麦蚜和龟纹瓢虫的的半变异函数均为指数模型,呈聚集分布。Block Kriging空间插值分析,得到二维空间插值模拟图,较好反映了麦蚜和龟纹瓢虫种群在麦田的时空分布。【结论】麦田生态系统中麦蚜与其天敌——龟纹瓢虫在时空分布上具有较强的相似性。同时研究结果提示,评价自然天敌对害虫的控害功能时,要综合分析两者的数量关系和空间分布关系及其控害功能的协同作用,才能提出可靠的防治策略。     

不同景观结构下麦田地面甲虫和蜘蛛物种多样性及优势种分布的时空动态

【目的】揭示在不同景观结构下,麦田地面甲虫和蜘蛛物种多样性及优势种分布的时空动态,为实际小麦生产过程和害虫种群控制提供理论上的依据和方法。【方法】采用地面陷阱法调查,对不同景观结构下地表甲虫和蜘蛛物种多样性的田间分布和时间动态进行分析与模拟。【结果】所得银川平原4种不同景观结构下麦田生境中地表甲虫和蜘蛛17科,92种,计5 683头。蜘蛛和步甲分别占总个体数的15.61%、31.15%。对不同景观结构下地表甲虫和蜘蛛物种多样性的田间分布和时间动态进行分析与模拟,得出:复杂景观结构较简单景观结构的调查区,表现出明显较高的物种多样性。步甲在5月上旬多样性指数最大,随后出现先降后升的趋势;蜘蛛在6月下旬多样性指数最大,之后一直呈现下降趋势。其原因主要是灌水影响了部分地表甲虫和蜘蛛活动所致。【结论】通过绘制了毛青步甲Chlaenius pallipes和星豹蛛Pardosa astrigera种群密度的田间分布图,可看出复杂景观结构调查区较简单景观结构调查区二者种群数量明显较高。     

区域农田景观格局对麦蚜种群数量的影响

明确农田景观格局对麦田蚜虫种群的影响,是开展区域性害虫生态调控的重要理论依据之一。以区域性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蚜虫种群数据,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个方面分析了区域农田景观格局对麦田蚜虫种群的影响。结果表明,蚜虫种群的数量与草地的平均斑块面积和最大斑块指数显著正相关,与县域的平均几何最邻近距离和面积加权平均斑块面积显著负相关,与耕地的面积加权平均斑块面积显著负相关,与耕地的斑块密度显著正相关。草地斑块面积的增大、区域景观与耕地的破碎化、区域景观的聚集会促进蚜虫种群数量的增加。使用草地的斑块面积和最大斑块指数、区域景观的平均几何最邻近距离可以预测蚜虫种群的发生量。非作物生境草地的斑块面积、耕地的破碎化、区域景观的空间分布及破碎化是影响麦田蚜虫种群发生的重要景观因素。     

黄河下游农业景观中不同生境类型地表节肢动物优势类群

生物多样性是连接景观异质性与生态系统服务的桥梁。在区域尺度上,以指示类群代替地表节肢动物类群,可以有效的开展生物多样性的保护工作。但是,在黄河下游农业景观中,关于地表节肢动物指示类群的研究报道较少。以黄河下游农业景观中4种生境类型(农田、林地、树篱和沟渠)为研究样地,且在农田生境中划分不同尺度(尺度1,3.6 hm2、尺度2,14 hm2和尺度3,28 hm2),通过调查不同生境中地表节肢动物分布及其多样性,结合地表节肢动物优势类群的辨识,分析了优势类群和地表节肢动物多样性的相关性,确定了研究区内地表节肢动物多样性的指示类群。结果发现:研究区内地表节肢动物优势类群为膜翅目、鞘翅目和蜘蛛目。树篱和林地生境的地表节肢动物多样性指示类群是膜翅目,农田生境中地表节肢动物多样性指示类群是鞘翅目,沟渠生境中地表节肢动物多样性指示类群是蜘蛛目。农田生境中优势类群间无相关性(P0.05);林地、沟渠和树篱生境中鞘翅目与蜘蛛目之间存在显著正相关(P0.01);林地和沟渠生境中,鞘翅目与膜翅目之间存在正相关(P0.05)。在农田生境中优势类群之间的相关性存在尺度依赖性,随着空间尺度的增大,相关性有一定的增强。在尺度2和尺度3上膜翅目和鞘翅目均存在正相关(P0.05),并且随着尺度增加而呈上升趋势。研究表明,黄河下游农业景观中不同生境类型地表节肢动物多样性的指示类群差别较大,其中树篱和林地生境具有较高的相似性,而农田生境地表节肢动物优势类群相关性存在尺度依赖性。在黄河下游农业景观中,以优势类群多样性代替地表节肢动物类群的多样性,可以在条件不足、时间紧迫的情况下更加有效的开展生物多样性的保护工作。     

瓢虫种群对棉花-玉米农田景观格局的响应

本文研究了华北棉花-玉米农田景观格局中龟纹瓢虫Propylaea japonica(Thunberg)和异色瓢虫Harmonia axyridis(Pallas)种群动态,发现农田景观格局中作物类型(棉花与玉米)对两种瓢虫种群密度动态有显著的影响,两种天敌瓢虫都趋向在玉米斑块上栖息。两种瓢虫在棉花斑块上呈现出时间分化,其中龟纹瓢虫在棉花种植的前中期种群密度较大,后期较少;而异色瓢虫在棉花前中期种群密度较少,后期较多,表明农田景观中种植玉米有利于增强瓢虫对棉花害虫的控制作用。进一步的研究表明,农田景观系统中玉米斑块所占的面积比对龟纹瓢虫和异色瓢虫种群密度均产生显著影响。这说明在农田景观系统中开展区域性生态调控的时候,需要考虑到各类斑块组合的面积比例,从而有利于增强多种天敌昆虫的协调控害作用。     

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.     

Predator–prey interactions in a ladybeetle–aphid system depend on spatial scale

The outcome of species interactions may manifest differently at different spatial scales; therefore, our interpretation of observed interactions will depend on the scale at which observations are made. For example, in ladybeetle–aphid systems, the results from small‐scale cage experiments usually cannot be extrapolated to landscape‐scale field observations. To understand how ladybeetle–aphid interactions change across spatial scales, we evaluated predator–prey interactions in an experimental system. The experimental habitat consisted of 81 potted plants and was manipulated to facilitate analysis across four spatial scales. We also simulated a spatially explicit metacommunity model parallel to the experiment. In the experiment, we found that the negative effect of ladybeetles on aphids decreased with increasing spatial scales. This pattern can be explained by ladybeetles strongly suppressing aphids at small scales, but not colonizing distant patches fast enough to suppress aphids at larger scales. In the experiment, the positive effects of aphids on ladybeetles were strongest at three‐plant scale. In a model scenario where predators did not have demographic dynamics, we found, consistent with the experiment, that both the effects of ladybeetles on aphids and the effects of aphids on ladybeetles decreased with increasing spatial scales. These patterns suggest that dispersal was the primary cause of ladybeetle population dynamics in our experiment: aphids increased ladybeetle numbers at smaller scales because ladybeetles stayed in a patch longer and performed area‐restricted searches after encountering aphids; these behaviors did not affect ladybeetle numbers at larger spatial scales. The parallel experimental and model results illustrate how predator–prey interactions can change across spatial scales, suggesting that our interpretation of observed predator–prey dynamics would differ if observations were made at different scales. This study demonstrates how studying ecological interactions at a range of scales can help link the results of small‐scale ecological experiments to landscape‐scale ecological problems.     

Seasonal differences in population‐, ensemble‐ and community‐level responses of bats to landscape structure in Amazonia

The amount (composition) and spatial arrangement (configuration) of forest patches in fragmented landscapes influence the accessibility, as well as the abundance and diversity of resources available to bats. Moreover, tropical fruit and insect abundance differ seasonally in response to changes in precipitation, and many bats in the family Phyllostomidae employ seasonal reproductive strategies. Because reproductive activities involve constraints on time and energy as well as increased nutritional demands, foraging behavior and home range size may differ between wet and dry seasons. Nonetheless, seasonal variation in response to landscape structure by bats has not been examined previously. Consequently, population‐, ensemble‐ and assemblage‐level responses of phyllostomids to landscape composition and configuration were quantified separately during the wet and dry season at three circular focal scales (1, 3 and 5 km radii) for 14 sites in fragmented lowland Amazon forest. Responses to landscape characteristics were scale‐dependent, species‐specific, and seasonal. Abundances of frugivores responded to landscape composition in the dry season and to landscape configuration in the wet season. Conversely, abundances of animalivores responded to landscape configuration in the dry season and to landscape composition in the wet season. Divergent responses to landscape structure between seasons suggest that variation in resource abundance and diversity play a significant role in structuring population‐, ensemble‐ and assemblage‐level patterns. As such, considerations of the effects of dietary flexibility and reproductive constraints on foraging strategies and habitat use may be important when designing management plans that successfully promote long‐term persistence of biodiversity in fragmented landscapes.     

海南北部蔬菜生境天敌瓢虫多样性研究

为了解海南北部主要蔬菜作物生境天敌瓢虫的群落结构和物种多样性，本文利用黄板诱集法于2019年对茄子、豇豆和辣椒生境的天敌瓢虫种类和数量进行了调查研究。结果表明，3种作物生境共采集获得瓢虫标本4 993头，隶属于16个属28种，其群落多样性从高到低依次为茄子、辣椒和豇豆生境，其中茄子生境拥有28种，辣椒和豇豆生境分别为10种和11种。茄子生境的多样性指数、丰富度指数、优势度指数和均匀度指数均为最高，分别为1.58、3.40、0.27和0.47，明显高于豇豆和辣椒生境。无论哪种生境，六斑月瓢虫Cheilomenes sexmaculata (Fabricius)和龟纹瓢虫Propylae japonica (Thunberg)均是优势种类，分别占整个瓢虫数量相对多度的45.26%和19.67%。两种优势瓢虫时序变化特征表明，瓢虫数量呈多峰型分布，5月中下旬至7月上旬这一时段总体发生数量较低。研究结果可为海南北部蔬菜作物生境天敌瓢虫保护和利用提供理论依据。     

Horizontal Variation of Bacterioplankton in the Baltic Sea

Variations in the thymidine incorporation rate, bacterial abundance, and mean cell volumes in the surface water (0.5 m) of the Baltic Sea in spring and summer were compared in studies with different spatial scales (570 nautical miles [nmi] [ca. 1056 km], 220 nmi [ca. 407 km], 24 nmi [ca. 44 km], 12 nmi [ca. 22 km], and 200 m). The objective of the comparison was to investigate whether a single sample taken at one sampling point is representative enough for researchers to make generalizations about a larger water area. Bacterioplankton variation was connected more to seasonal characteristics than to the spatial scale of sampling. Variation was greater and more random in spring than in summer. The state variables (bacterial abundance and mean cell volume) varied less than the rate variable (thymidine incorporation). The results suggest that the sampling design for bacterioplankton studies in northern temperate seas should be planned primarily according to the season and that more stress should be put on rate variable measurements than on those of state variables.     

Influence of Landscape Diversity and Composition on the Parasitism of Cotton Bollworm Eggs in Maize

We deployed >50,000 Helicoverpa armigera eggs in maize fields to assess the rate of parasitism by Trichogramma chilonis across 33 sites during a three-year span (2012–2014) in northern China. Subsequently, we used a partial least squares (PLS) regression approach to assess the relationship of landscape diversity with composition and parasitism potential. The parasitism rate of H. armigera eggs by T. chilonis ranged from 0–25.8%, with a mean value of 5.6%. Landscape diversity greatly enhanced parasitism at all four different spatial scales (0.5, 1.0, 1.5 and 2.0 km radius). Both the proportion of arable area and the total planting area of two major crops (cotton and maize) had a negative correlation to the parasitism rate at each scale, whereas parasitism was positively correlated to the proportion of host crops of H. armigera other than cotton and maize at the 0.5 to 2.0 km radius scales as well as to that of non-crop habitat at the 0.5 and 1.0 km radius scales. The study indicated that maintaining landscape diversity provided an important biocontrol service by limiting H. armigera through the egg parasitoid T. chilonis, whereas rapid agricultural intensification would greatly reduce the presence and parasitism of T. chilonis in China.     

Effects of landscape complexity and stand factors on arthropod communities in poplar forests

The arthropod communities are influenced by both local conditions and features of the surrounding landscape. Landscape complexity and stand factors may both influence arthropod communities in poplar forests, but the multiscale effects of these factors on poplar defoliators and natural enemies are still poorly understood. We collected poplar arthropods at 30 sampling sites within five forest landscapes in Xinjiang, China, and assessed whether landscape complexity and stand factors influence species abundance and diversity of poplar arthropods. Landscape complexity was quantified by several independent metrics of landscape composition, configuration, and connectivity at three spatial scales. We also determined the most powerful explanatory variables and the scale effect of each arthropod. Results found that landscape complexity and stand factors had different effects on different poplar arthropod communities. Landscape complexity promoted natural enemies at different spatial scales, but it inhibited the population of poplar defoliators at the scale of 200 m. Specifically, the abundance and diversity of all defoliators decreased with increasing proportion of nonhost plants. Landscape diversity only had a negative effect on defoliator abundance. The shape complexity of habitat patches increased the abundance of carabid beetles but reduced the abundance of green leafhoppers and migratory locusts. The abundance and diversity of predators increased with increasing structural connectivity of forest landscape. Additionally, both the abundance and diversity of all defoliators were positively correlated with the average height of herbaceous plants. Diversity of all defoliators increased with increasing size of host trees. The distance from sampling site to the nearest village positively influenced the abundance and diversity of all predators. Arthropod abundance and diversity in poplar forests were driven by stand factors and landscape complexity. Therefore, maintaining complex shape and structural connectivity of habitat patches and keeping poplar stands away from the village are crucial for management of forest landscape to enhance natural enemies. And in order to reduce the abundance of defoliators in poplar forest, the diversity of surrounding habitat types should be promoted within 200 m radii.     

The landscape context of cereal aphid-parasitoid interactions

Analyses at multiple spatial scales may show how important ecosystem services such as biological control are determined by processes acting on the landscape scale. We examined cereal aphid-parasitoid interactions in wheat fields in agricultural landscapes differing in structural complexity (32-100% arable land). Complex landscapes were associated with increased aphid mortality resulting from parasitism, but also with higher aphid colonization, thereby counterbalancing possible biological control by parasitoids and lastly resulting in similar aphid densities across landscapes. Thus, undisturbed perennial habitats appeared to enhance both pests and natural enemies. Analyses at multiple spatial scales (landscape sectors of 0.5-6 km diameter) showed that correlations between parasitism and percentage of arable land were significant at scales of 0.5-2 km, whereas aphid densities responded to percentage of arable land at scales of 1-6 km diameter. Hence, the higher trophic level populations appeared to be determined by smaller landscape sectors owing to dispersal limitation, showing the 'functional spatial scale' for species-specific landscape management.     

Spatial pattern of the threatened epiphytic bryophyte Neckera pennata at two scales in a fragmented boreal forest

The spatial pattern and occurrence of a threatened bryophyte, Neckera pennata. were studied in relation to the abundance and pattern of suitable substrate trees at two spatial scales: 1) in a 4 x 4 km fraction of fragmented, mostly managed southern boreal forest landscape, and 2) in an old-growth forest stand within this landscape, with abundant occurrence of suitable habitats. To explore in detail the spatial clustering of N. pennata at the forest stand scale, we applied a second order point process analysis based on the Ripley's K-function for binary point patterns, Neckera pennata proved to be a rare species in the studied landscape: it was found only on 31 Populus tremula trees. At the landscape scale, the distribution of the species was highly aggregated: the species occurred only within a 60 ha old-growth forest patch in the whole area. At the forest stand scale, N. pennata proved to be a widespread, randomly distributed species without any tendency towards aggregation. It was found on 19 Populus trees, which was only 1.5% of the total 1253 potential substrate trees within the inventory area. The species showed a statistically significant preference towards large trees. The future of the species in the study area is unclear due to 1) the very low population density and 2) the lack of regeneration of Populus within the protected old-growth forest area hosting the remaining population.     

Landscape heterogeneity at multiple spatial scales enhances spider species richness in an agricultural landscape

Landscape supplementation, which enhances densities of organisms by combination of different landscape elements, is likely common in heterogeneous landscapes, but its prevalence and effects on species richness have been little explored. Using grassland-dwelling spiders in an agricultural landscape, we postulated that richness and abundances of major constituent species are both highest in intermediate mixtures of forests and paddy fields, and that this effect derives from multi-scale landscape heterogeneity. We collected spiders in 35 grasslands in an agricultural landscape in Japan and determined how species richness and abundances of major species related to local and landscape factors across different spatial scales. We used a generalized linear model to fit data, created all possible combinations of variables at 15 spatial scales, and then explored the best models using Akaike's information criterion. Species richness showed a hump-shaped pattern in relation to surrounding forest cover, and the spatial scale determining this relationship was a 300–500-m radius around the study sites. Local variables were of minor importance for species richness. Abundances of major species also exhibited a hump-shaped pattern when plotted against forest cover. Thus, a combination of paddy fields and forests is important for enhancement of grassland spider species richness and abundance, suggesting habitat supplementation. The effective spatial scales determining abundances varied, ranging from 200 to >1000 m, probably representing different dispersal abilities. Landscape compositional heterogeneity at multiple spatial scales may be thus crucial for the maintenance of species diversity.