Stability and synchrony across ecological hierarchies in heterogeneous metacommunities: linking theory to data

Understanding stability across ecological hierarchies is critical for landscape management in a changing world. Recent studies showed that synchrony among lower‐level components is key to scaling temporal stability across two hierarchical levels, whether spatial or organizational. But an extended framework that integrates both spatial scale and organizational level simultaneously is required to clarify the sources of ecosystem stability at large scales. However, such an extension is far from trivial when taking into account the spatial heterogeneities in real‐world ecosystems. In this paper, we develop a partitioning framework that bridges variability and synchrony measures across spatial scales and organizational levels in heterogeneous metacommunities. In this framework, metacommunity variability is expressed as the product of local‐scale population variability and two synchrony indices that capture the temporal coherence across species and space, respectively. We develop an R function ‘var.partition’ and apply it to five types of desert plant communities to illustrate our framework and test how diversity shapes synchrony and variability at different hierarchical levels. As the observation scale increased from local populations to metacommunities, the temporal variability of plant productivity was reduced mainly by factors that decreased species synchrony. Species synchrony decreased from local to regional scales, and spatial synchrony decreased from species to community levels. Local and regional species diversity were key factors that reduced species synchrony at the two scales. Moreover, beta diversity contributed to decreasing spatial synchrony among communities. We conclude that our new framework offers a valuable toolbox for future empirical studies to disentangle the mechanisms and pathways by which ecological factors influence stability at large scales.     

北京昌平区景观复杂度和局地管理对苹果园传粉蜂多样性的影响

传粉蜂为作物生产和粮食安全提供重要的生态系统服务。随着农业经济的不断发展,土地利用强度加剧,大量自然或半自然生境已经转变为农业用地。景观均质化和集约化管理导致野生蜂多样性下降,从而威胁到农业可持续生产。本研究以北京市昌平区苹果园为对象,探究景观复杂度(半自然生境比例)、局地管理强度(地表开花植物多样性和土壤全氮)及其交互作用对传粉蜂多样性的影响。结果表明: 共捕获传粉蜂8642头,其中人工蜂5125头,野生蜂分属5科14属49种3517头。传粉蜂多样性对景观复杂度和局地管理强度响应的最优尺度在500 m。样点半径500 m范围内,总传粉蜂和野生蜂多度随周围半自然生境增加均呈显著上升趋势。景观复杂度与开花植物多样性的交互作用对总传粉蜂和野生蜂物种丰富度有显著影响。当景观复杂度较低时(≤29.9%),总传粉蜂和野生蜂物种丰富度与开花植物多样性呈显著正相关;而当景观复杂度较高时(>29.9%),总传粉蜂和野生蜂物种丰富度与开花植物多样性呈显著负相关。此外,人工蜂多度随果园内局地开花植物多样性和土壤全氮增加呈显著升高趋势。土壤全氮与开花植物多样性的交互作用对人工蜂多度有显著影响。当土壤全氮含量较低时(≤1.9 g·kg^-1),人工蜂多度与开花植物多样性呈显著正相关;而当土壤全氮含量较高时(>1.9 g·kg^-1),人工蜂多度与开花植物多样性呈显著负相关。农业景观中半自然生境比例的增加有利于提高野生蜂多度,而地表开花植物多样性可以促进传粉蜂多样性,但是受到景观尺度(半自然生境比例)和局地尺度(氮肥施用)的影响。因此,农业景观中野生蜂多样性的维持需要综合考虑多尺度因素来制定保护策略。尽可能保留更高比例的耕地仍然是生产的长期需求,而保持中等景观复杂度,增加地表开花植物多样性,减少氮肥施用量将是促进苹果园传粉蜂多样性的有效方式。     

Solar radiation and functional traits explain the decline of forest primary productivity along a tropical elevation gradient

One of the major challenges in ecology is to understand how ecosystems respond to changes in environmental conditions, and how taxonomic and functional diversity mediate these changes. In this study, we use a trait‐spectra and individual‐based model, to analyse variation in forest primary productivity along a 3.3 km elevation gradient in the Amazon‐Andes. The model accurately predicted the magnitude and trends in forest productivity with elevation, with solar radiation and plant functional traits (leaf dry mass per area, leaf nitrogen and phosphorus concentration, and wood density) collectively accounting for productivity variation. Remarkably, explicit representation of temperature variation with elevation was not required to achieve accurate predictions of forest productivity, as trait variation driven by species turnover appears to capture the effect of temperature. Our semi‐mechanistic model suggests that spatial variation in traits can potentially be used to estimate spatial variation in productivity at the landscape scale.     

Biodiversity–productivity relationship in ponds: Community and metacommunity patterns along time and environmental gradients

Primary production correlates with diversity in various ways. These patterns may result from the interaction of various mechanisms related to the environmental context and the spatial and temporal scale of analysis. However, empirical evidence on diversity‐productivity patterns typically considers single temporal and spatial scales, and does not include the effect of environmental variables. In a metacommunity of macrophytes in ephemeral ponds, we analysed the diversity‐productivity relationship patterns in the field, the importance of the environmental variables of pond size and heterogeneity on such relationship, and the variation of these patterns at local (community level) and landscape scales (metacommunity level) across 52 ponds on twelve occasions, over five years (2005–2009). Combining all sampling dates, there were 377 ponds and 1954 sample‐unit observations. Vegetation biomass was used as a proxy for productivity, and biodiversity was represented by species richness, evenness, and their interaction. Environmental variables comprised pond area, depth and internal heterogeneity. Productivity and species richness were not directly related at the metacommunity level, and were positively related at the community level. Taking environmental variables into account revealed positive species richness‐productivity relationships at the metacommunity level and positive quadratic relationships at the community level. Productivity showed both positive and negative linear and nonlinear relationships with the size and heterogeneity of ponds. We found a weak relationship between productivity and evenness. The identity of variables associated with productivity changed between spatial scales and through time. The pattern of relationships between productivity and diversity depends on spatial scale and environmental context, and changes idiosyncratically through time within the same ecosystem. Thus, the diversity‐productivity relationship is not only a property of the study system, but also a consequence of environmental variations and the temporal and spatial scale of analysis.     

南京东郊城市湿地水质多尺度空间分异

对南京市东郊的17个城市湿地进行水质监测和植物群落调查,从生态系统、集水区和景观功能区3个尺度揭示城市湿地水质的分异特征.结果表明:湿地类型和植物群落特征是生态系统尺度上影响湿地水质的主要因素;原生湿地的水质普遍优于改造湿地,而改造湿地的水质则普遍优于次生湿地;湿地植物的盖度越大,水质越好.土地利用类型是集水区尺度上影响湿地水质空间分异的主要因素,建设用地对流域湿地营养盐影响程度较大,而林地对溶解氧影响程度较大.人类活动是功能区尺度上影响湿地水质空间分异的主要因素,而其对不同水质指标的影响程度不同.     

Spatial scaling of arbuscular mycorrhizal fungal diversity is affected by farming practice

Evidence suggests that microbial communities show patterns of spatial scaling which can be driven by geographical distance and environmental heterogeneity. Here we demonstrate that human management can have a major impact on microbial distribution patterns at both the local and landscape scale. Mycorrhizal fungi are vital components of terrestrial ecosystems, forming a mutualistic symbiosis with plant roots which has a major impact on above ground ecology and productivity. We used contrasting agricultural systems to investigate the spatial scaling of the most widespread mycorrhizal fungus group, the arbuscular mycorrhizal fungi (AMF). Using multiple sampling sites with a maximum separation of 250 km we describe for the first time the roles which land management, environmental heterogeneity and geographical distance play in determining spatial patterns of microbial distribution. Analysis of AMF taxa–area relationships at each sampling site revealed that AMF diversity and spatial turnover was greater under organic relative to conventional farm management. At the regional scale (250 km) distance–decay analyses showed that there was significant change in AMF community composition with distance, and that this was greater under organic relative to conventional management. Environmental heterogeneity was found to be the major factor determining turnover of AMF taxa at the landscape scale. Overall we demonstrate that human management can play a key role in determining the turnover of microbial communities at both the local and regional scales.     

Pollination and other ecosystem services produced by mobile organisms: a conceptual framework for the effects of land-use change

Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60–90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.     

外来植物入侵对生态系统生产力的影响研究综述

外来植物入侵已成为全球尺度生态系统面临的一个主要环境问题。分析和总结了外来植物入侵对于生态系统生物量、生产力和碳固定的影响以及主要的影响因素,多数研究结果显示外来植物入侵会增加生态系统生物量和生产力,并进而增加生态系统对于二氧化碳气体的固定,但也有些研究结果得出相反的结论;另外,造成增加或减少结果的成因也是复杂的。系统全面地研究外来植物入侵,特别是考虑到空间上异质性和入侵的时间尺度,在研究外来植物入侵对生态系统生物量、生产力以及碳固定的影响方面尤为重要。     

Plant species richness and productivity determine the diversity of soil fungal guilds in temperate coniferous forest and bog habitats

Fungi have important roles as decomposers, mycorrhizal root symbionts and pathogens in forest ecosystems, but there is limited information about their diversity and composition at the landscape scale. This work aimed to disentangle the factors underlying fungal richness and composition along the landscape‐scale moisture, organic matter and productivity gradients. Using high‐throughput sequencing, we identified soil fungi from 54 low‐productivity Pinus sylvestris‐dominated plots across three study areas in Estonia and determined the main predictors of fungal richness based on edaphic, floristic and spatial variables. Fungal richness displayed unimodal relationship with organic matter and deduced soil moisture. Plant richness and productivity constituted the key predictors for taxonomic richness of functional guilds. Composition of fungi and the main ectomycorrhizal fungal lineages and hyphal exploration types was segregated by moisture availability and soil nitrogen. We conclude that plant productivity and diversity determine the richness and proportion of most functional groups of soil fungi in low‐productive pine forests on a landscape scale. Adjacent stands of pine forest may differ greatly in the dominance of functional guilds that have marked effects on soil carbon and nitrogen cycling in these forest ecosystems.     

Meta-ecosystems: a theoretical framework for a spatial ecosystem ecology

This contribution proposes the meta‐ecosystem concept as a natural extension of the metapopulation and metacommunity concepts. A meta‐ecosystem is defined as a set of ecosystems connected by spatial flows of energy, materials and organisms across ecosystem boundaries. This concept provides a powerful theoretical tool to understand the emergent properties that arise from spatial coupling of local ecosystems, such as global source–sink constraints, diversity–productivity patterns, stabilization of ecosystem processes and indirect interactions at landscape or regional scales. The meta‐ecosystem perspective thereby has the potential to integrate the perspectives of community and landscape ecology, to provide novel fundamental insights into the dynamics and functioning of ecosystems from local to global scales, and to increase our ability to predict the consequences of land‐use changes on biodiversity and the provision of ecosystem services to human societies.     

Functional spatial scale of community composition change in response to windthrow disturbance in a deciduous temperate forest

Community dynamics in local habitats are affected by landscape characteristics such as the area and connectivity of surrounding habitats at a functional spatial scale where the community responds to landscape structure. However, the functional spatial scale at which community composition is affected by landscape structure has never been explored. We assessed the functional spatial scales of composition change in birds and in three types of arthropod communities (canopy, forest-floor and flying ones) with regard to landscape heterogeneity resulting from a large typhoon in a temperate forest of Japan. We examined the effects of tree-fall disturbance on the communities at various spatial scales, with special attention to compositional evenness. The spatial scale of the best-fitting model, which was selected from models fitted to the disturbance area at stepwise spatial scales, was interpreted as the community-specific functional spatial scale. The composition of all communities studied was all significantly dependent on gap area. The functional spatial scale was highest in birds (370 m in radius), intermediate in flying arthropods (90 m) and lowest in canopy and forest-floor arthropods (10 m). This result may reflect typical dispersal ability and the spatial range of resource use in the community. Compositional changes in each community were consistent with theory regarding traits and responses of component taxa, although the enhancement of evenness was observed only in the arthropod communities. These results imply that management and reserve selection based on functional spatial scales can be effective in the conservation of biodiversity and ecosystem services at the community level.     

Productivity–species richness relationship changes from unimodal to positive linear with increasing spatial scale in the Inner Mongolia steppe

Productivity–species diversity relationships have been a controversial research topic in ecology with scale believed to be among the main reasons for discovering different relationships. We collected data on species diversity (richness) and productivity (peak above-ground biomass) of the Stipa breviflora association in the Inner Mongolia grassland to examine spatial scale dependency and possible underlying mechanisms responsible for the relationships found. One local and seven different landscape scales (the first level corresponds in extent to a 100 × 100 km area, which is increased consecutively by 100 km resulting in the 700 × 700 km area at the highest level) were considered. We found that: (1) unimodal relationships dominated the local scale, but this varied depending on the position along successional gradients; (2) a positive linear relationship was common at larger spatial scales; (3) biotic processes were the most likely primary factor underlying local scale unimodal relationships, but environmental heterogeneity (precipitation patterns) was the main determinant of relationships found at larger spatial scales; (4) our study contributed to other empirical evidence and predictions of theoretical models regarding scale dependency of productivity–species richness relationships; (5) while earlier research demonstrated positive linear species richness–productivity relationships across a number of ecological scales in the Inner Mongolia steppe, our study specifically tested a spectrum of geographical scales to confirm the scale-dependency of this relationship. Lastly, our study emphasized the critical role played by precipitation patterns in controlling biodiversity and grassland ecosystem functioning, which maintains the relatively high level of biodiversity and stable ecosystem processes.     

Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences

In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species‐richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co‐existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation. In contrast to the diversifying effects of local disturbances, productivity‐related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro‐evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats. During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co‐existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.     

多尺度生态系统服务的权衡关系及其对景观配置的响应——以湖北省为例

多种生态系统服务相互关系及其对景观配置响应的多尺度分析一直是生态系统服务研究的热点问题,对于厘清生态系统服务尺度效应和实现区域分级管理具有重要的意义。以湖北省为研究区域,采用遥感、土地利用类型、水文、气象、土壤等多元数据,从5 km×5 km、10 km×10 km、15 km×15 km、20 km×20 km、25 km×25 km、县域和市域等7个递增尺度分析4种生态系统服务(水源供给、碳固持、粮食生产、水质净化)对景观配置的响应关系。结果表明：(1)4种生态系统服务的空间分布具有较强的异质性,粮食生产服务的高值区与水质净化服务的低值区主要分布于湖北省中部和东部的耕地集中区域,碳固持与水源供给呈现相反的空间特征。(2)生态系统服务权衡关系的作用强度随尺度增加而减弱,而协同关系呈增加趋势;同时服务间相互作用强度的变化大于作用方向的变化。(3)景观配置对生态系统服务的影响程度在县域和市域尺度上大于格网尺度。结果反映了受尺度变化中“削峰填谷”过程的影响,景观配置对生态系统服务影响的大小和正负效应具有尺度差异性。研究结果可为区域生态系统服务分级管理和景观格局调控提供科学参考。     

Landscape structure effects on forest plant diversity at local scale: Exploring the role of spatial extent

Since landscape attributes show different patterns at different spatial extents, it is fundamental to identify how the relation between landscape structure and plant species diversity at local scale varies with scale. Then, it is fundamental to assess the appropriate extent at which landscape factors affect plant species richness at the local scale. To investigate this relation, data on plant species richness of forest communities at plot scale were extracted from a large data set and landscape metrics were calculated around the same plots for a range of extents (250–3000 m). Then, multiple regression models and variance partitioning techniques were applied to assess the amount of variance explained by the landscape metrics on plant species richness for a range of extents. In general, we found that increasing extent of the surrounding landscape analyzed, improved the strength of relationship between the landscape metrics and the properties of plant communities at plot scale. The medium-large extent was most informative as it combined a decent total variance explained with high variance explained by the pure fractions of complexity, fragmentation and disturbance and the minimum of collinearity. In conclusion, we found that it is possible and beneficial to identify a specific extent, where the redundancy in the predictor variables is minimized and the explanatory power of the pure fractions (or single groups) maximized, when examining landscape structure effects on local plant species richness.     

An integrated approach to identify distribution of Oncomelania hupensis, the intermediate host of Schistosoma japonicum, in a mountainous region in China

The aim of this study is to better understand ecological variability related to the distribution of Oncomelania hupensis, the snail intermediate host of Schistosoma japonicum, and predict the spatial distribution of O. hupensis at the local scale in order to develop a more effective control strategy for schistosomiasis in the hilly and mountainous regions of China. A two-pronged approach was applied in this study consisting of a landscape pattern analysis complemented with Bayesian spatial modelling. The parasitological data were collected by cross-sectional surveys carried out in 11 villages in 2006 and mapped based on global positioning system (GPS) coordinates. Environmental surrogates and landscape metrics were derived from remotely-sensed images and land-cover/land-use classification data. Bayesian non-spatial and spatial models were applied to investigate the variation of snail density in relation to environmental surrogates and landscape metrics at the local scale. A Bayesian spatial model, validated by the deviance information criterion (DIC), was found to be the best-fitting model. The mean shape index (MSI) and Shannon's evenness indexes (SEI) were significantly associated with snail density. These findings suggest that decreasing the heterogeneity of the landscape can reduce snail density. A prediction maps were generated by the Bayesian model together with environmental surrogates and landscape metrics. In conclusion, the risk areas of snail distribution at the local scale can be identified using an integrated approach with landscape pattern analysis supported by remote sensing and GIS technologies, as well as Bayesian modelling.     

Local and landscape‐scale effects of an ant nest construction in an open dry forest of Uruguay

1. Ants modify soil properties via nest construction and by doing this modulate soil resources for other organisms. In this sense ants are recognised as ecosystem engineers. 2. In this framework, two less well‐studied issues are focused on: (i) the permanence of the effects of ant nests on plant communities after colonies have died, and (ii) the scaling up from patch to landscape‐scale effects. 3. The aim of the present study was to address these issues in the open dry forests of Uruguay, inhabited by the ant Atta vollenweideri Forel. The active and abandoned nests of this ant represent different and conspicuous patches (30–60 m²) in the landscape. 4. The soil concentration of sodium, a key element in the system, was substantially higher among active nests, and remains high during the early stages of abandoned nests. Woody species abundance, richness, and composition were affected at the patch scale, and simulation models suggested an increase in species richness at the landscape scale. 5. The present study highlights the importance of abandoned nests for plant‐species richness in the ecosystem engineer framework and the need to advance in an integrative approach to study both local and landscape effects of ant's nests.     

Local diversity of arable weeds increases with landscape complexity

Patterns of plant diversity are often related to local site conditions and to competitive interactions, but landscape context may also be important for local plant species richness. This is shown here by analysing the relationship between landscape complexity and local species richness of arable weeds in wheat fields. The fields were located in 18 landscapes characterised by a gradient in landscape complexity from structurally complex to structurally simple (39–94% arable land). We quantified local site conditions, field management intensity and landscape characteristics, and used principle component analyses to ordinate the environmental variables. The percentage of arable land was negatively correlated with perimeter–area ratio, habitat-type diversity and topographical heterogeneity, but landscape characteristics did not correlate with local site conditions and field management intensity. The number of plant species was mainly related to landscape characteristics and to a lesser extent to field management intensity (nitrogen fertilisation), whereas local soil characteristics did not contribute to the explanation of arable weed richness. In a geographic scale analysis using circular landscape sectors ranging from 1 km up to 5 km diameter, the predictive power of landscape complexity for local plant species richness was strongest at 2 km indicating a scale-dependent relationship between landscape context and plant species richness. Our results support the hypothesis that local plant species richness in arable fields is greatly influenced by processes operating at the landscape scale. Seed rain from ruderal source habitats and disturbed edges may be the most important underlying process.     

A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in‐field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in‐field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes.