Reconciling complexity with stability in naturally assembling food webs

Understanding how complex food webs assemble through time is fundamental both for ecological theory and for the development of sustainable strategies of ecosystem conservation and restoration. The build-up of complexity in communities is theoretically difficult, because in random-pattern models complexity leads to instability. There is growing evidence, however, that nonrandom patterns in the strengths of the interactions between predators and prey strongly enhance system stability. Here we show how such patterns explain stability in naturally assembling communities. We present two series of below-ground food webs along natural productivity gradients in vegetation successions. The complexity of the food webs increased along the gradients. The stability of the food webs was captured by measuring the weight of feedback loops of three interacting 'species' locked in omnivory. Low predator-prey biomass ratios in these omnivorous loops were shown to have a crucial role in preserving stability as productivity and complexity increased during succession. Our results show the build-up of food-web complexity in natural productivity gradients and pin down the feedback loops that govern the stability of whole webs. They show that it is the heaviest three-link feedback loop in a network of predator-prey effects that limits its stability. Because the weight of these feedback loops is kept relatively low by the biomass build-up in the successional process, complexity does not lead to instability.     

Biodiversity and ecosystem stability in a decade-long grassland experiment

Human-driven ecosystem simplification has highlighted questions about how the number of species in an ecosystem influences its functioning. Although biodiversity is now known to affect ecosystem productivity, its effects on stability are debated. Here we present a long-term experimental field test of the diversity-stability hypothesis. During a decade of data collection in an experiment that directly controlled the number of perennial prairie species, growing-season climate varied considerably, causing year-to-year variation in abundances of plant species and in ecosystem productivity. We found that greater numbers of plant species led to greater temporal stability of ecosystem annual aboveground plant production. In particular, the decadal temporal stability of the ecosystem, whether measured with intervals of two, five or ten years, was significantly greater at higher plant diversity and tended to increase as plots matured. Ecosystem stability was also positively dependent on root mass, which is a measure of perenniating biomass. Temporal stability of the ecosystem increased with diversity, despite a lower temporal stability of individual species, because of both portfolio (statistical averaging) and overyielding effects. However, we found no evidence of a covariance effect. Our results indicate that the reliable, efficient and sustainable supply of some foods (for example, livestock fodder), biofuels and ecosystem services can be enhanced by the use of biodiversity.     

Ecosystem size determines food-chain length in lakes

Food-chain length is an important characteristic of ecological communities: it influences community structure, ecosystem functions and contaminant concentrations in top predators. Since Elton first noted that food-chain length was variable among natural systems, ecologists have considered many explanatory hypotheses, but few are supported by empirical evidence. Here we test three hypotheses that predict food-chain length to be determined by productivity alone (productivity hypothesis), ecosystem size alone (ecosystem-size hypothesis) or a combination of productivity and ecosystem size (productive-space hypothesis). The productivity and productive-space hypotheses propose that food-chain length should increase with increasing resource availability; however, the productivity hypothesis does not include ecosystem size as a determinant of resource availability. The ecosystem-size hypothesis is based on the relationship between ecosystem size and species diversity, habitat availability and habitat heterogeneity. We find that food-chain length increases with ecosystem size, but that the length of the food chain is not related to productivity. Our results support the hypothesis that ecosystem size, and not resource availability, determines food-chain length in these natural ecosystems.     

Consistent patterns and the idiosyncratic effects of biodiversity in marine ecosystems

Revealing the consequences of species extinctions for ecosystem function has been a chief research goal and has been accompanied by enthusiastic debate. Studies carried out predominantly in terrestrial grassland and soil ecosystems have demonstrated that as the number of species in assembled communities increases, so too do certain ecosystem processes, such as productivity, whereas others such as decomposition can remain unaffected. Diversity can influence aspects of ecosystem function, but questions remain as to how generic the patterns observed are, and whether they are the product of diversity, as such, or of the functional roles and traits that characterize species in ecological systems. Here we demonstrate variable diversity effects for species representative of marine coastal systems at both global and regional scales. We provide evidence for an increase in complementary resource use as diversity increases and show strong evidence for diversity effects in naturally assembled communities at a regional scale. The variability among individual species responses is consistent with a positive but idiosyncratic pattern of ecosystem function with increased diversity.     

Effects of species and functional group loss on island ecosystem properties

Considerable recent attention has focused on predicting how the losses of species and functional groups influence ecosystem properties, but the extent to which these effects vary among ecosystems remains poorly understood. Island systems have considerable scope for studying how biotic and abiotic factors influence processes in different ecosystems, because they enable the simultaneous study of large numbers of independent replicate systems at ecologically meaningful spatial scales. We studied a group of 30 islands in northern Sweden, for which island size determined disturbance history, and therefore vegetation successional stage and biotic and abiotic ecosystem properties. On each island we conducted a seven-year study that involved experimental removals of combinations of both plant functional groups and plant species. We show that although losses of functional groups and species often impaired key ecosystem processes, these effects were highly context-dependent and strongly influenced by island size. Our study provides evidence that the consequences of biotic loss for ecosystem functioning vary greatly among ecosystems and depend on the specific abiotic and biotic attributes of the system.     

Ecology: diversity and stability in plant communities

The relationship between species diversity and ecosystem stability is controversial. Tilman et al. analyse biomass patterns over a decade in a grassland experiment with artificial plant communities, and provide evidence for a positive relationship between the number of plant species and the temporal stability of the ecosystem. Here we use data from a long-term biodiversity experiment with plant communities that were not controlled by weeding in order to show that diverse systems can be both stable and unstable.     

Consumer versus resource control of species diversity and ecosystem functioning

A key question in ecology is which factors control species diversity in a community. Two largely separate groups of ecologists have emphasized the importance of productivity or resource supply, and consumers or physical disturbance, respectively. These variables show unimodal relationships with diversity when manipulated in isolation. Recent multivariate models, however, predict that these factors interact, such that the disturbance diversity relationship depends on productivity, and vice versa. We tested these models in marine food webs, using field manipulations of nutrient resources and consumer pressure on rocky shores of contrasting productivity. Here we show that the effects of consumers and nutrients on diversity consistently depend on each other, and that the direction of their effects and peak diversity shift between sites of low and high productivity. Factorial meta-analysis of published experiments confirms these results across widely varying aquatic communities. Furthermore, our experiments demonstrate that these patterns extend to important ecosystem functions such as carbon storage and nitrogen retention. This suggests that human impacts on nutrient supply and food-web structure have strong and interdependent effects on species diversity and ecosystem functioning, and must therefore be managed together.     

Ecosystem stability and compensatory effects in the Inner Mongolia grassland

Numerous studies have suggested that biodiversity reduces variability in ecosystem productivity through compensatory effects; that is, a species increases in its abundance in response to the reduction of another in a fluctuating environment. But this view has been challenged on several grounds. Because most studies have been based on artificially constructed grasslands with short duration, long-term studies of natural ecosystems are needed. On the basis of a 24-year study of the Inner Mongolia grassland, here we present three key findings. First, that January-July precipitation is the primary climatic factor causing fluctuations in community biomass production; second, that ecosystem stability (conversely related to variability in community biomass production) increases progressively along the hierarchy of organizational levels (that is, from species to functional group to whole community); and finally, that the community-level stability seems to arise from compensatory interactions among major components at both species and functional group levels. From a hierarchical perspective, our results corroborate some previous findings of compensatory effects. Undisturbed mature steppe ecosystems seem to culminate with high biodiversity, productivity and ecosystem stability concurrently. Because these relationships are correlational, further studies are necessary to verify the causation among these factors. Our study provides new insights for better management and restoration of the rapidly degrading Inner Mongolia grassland.     

Diversity and dispersal interactively affect predictability of ecosystem function

Theory and small-scale experiments predict that biodiversity losses can decrease the magnitude and stability of ecosystem services such as production and nutrient cycling. Most of this research, however, has been isolated from the immigration and emigration (dispersal) processes that create and maintain diversity in nature. As common anthropogenic drivers of biodiversity change--such as habitat fragmentation, species introductions and climate change--are mediated by these understudied processes, it is unclear how environmental degradation will affect ecosystem services. Here we tested the interactive effects of mobile grazer diversity and dispersal on the magnitude and stability of ecosystem properties in experimental seagrass communities that were either isolated or connected by dispersal corridors. We show that, contrary to theoretical predictions, increasing the number of mobile grazer species in these metacommunities increased the spatial and temporal variability of primary and secondary production. Moreover, allowing grazers to move among and select patches reduced diversity effects on production. Finally, effects of diversity on stability differed qualitatively between patch and metacommunity scales. Our results indicate that declining biodiversity and habitat fragmentation synergistically influence the predictability of ecosystem functioning.     

Diversity-dependent production can decrease the stability of ecosystem functioning

There is concern that species loss may adversely affect ecosystem functioning and stability. But although there is evidence that biodiversity loss can lead to reductions in biomass production, there is no direct evidence that biodiversity loss affects ecosystem resistance (ability to withstand perturbation) or resilience (recovery from perturbation). Yet theory, laboratory experiments and indirect experimental evidence strongly suggest that diversity and stability are related. Here we report results from a field experiment with factorially crossed perturbation and diversity manipulations. We simulated drought perturbation on constructed grassland ecosystems containing 1, 2, 4, 8 or 32 plant species. Under unperturbed conditions, the species-poor systems achieved lower biomass production than the species-rich systems. However, the species-poor systems were more resistant to perturbation than the species-rich systems. The species-poor systems also showed a larger initial resilience following perturbation, although the original relationship between diversity and productivity was fully restored after 1year. Our results confirm that biodiversity increases biomass production, but they also point to the fact that such diversity--production associations may lead to an inverse relationship between biodiversity and the stability of ecosystem functioning.     

Recuperation of nitrogen cycling in Amazonian forests following agricultural abandonment

Phosphorus (P) is generally considered the most common limiting nutrient for productivity of mature tropical lowland forests growing on highly weathered soils. It is often assumed that P limitation also applies to young tropical forests, but nitrogen (N) losses during land-use change may alter the stoichiometric balance of nutrient cycling processes. In the Amazon basin, about 16% of the original forest area has been cleared, and about 30-50% of cleared land is estimated now to be in some stage of secondary forest succession following agricultural abandonment. Here we use forest age chronosequences to demonstrate that young successional forests growing after agricultural abandonment on highly weathered lowland tropical soils exhibit conservative N-cycling properties much like those of N-limited forests on younger soils in temperate latitudes. As secondary succession progresses, N-cycling properties recover and the dominance of a conservative P cycle typical of mature lowland tropical forests re-emerges. These successional shifts in N:P cycling ratios with forest age provide a mechanistic explanation for initially lower and then gradually increasing soil emissions of the greenhouse gas nitrous oxide (N(2)O). The patterns of N and P cycling during secondary forest succession, demonstrated here over decadal timescales, are similar to N- and P-cycling patterns during primary succession as soils age over thousands and millions of years, thus revealing that N availability in terrestrial ecosystems is ephemeral and can be disrupted by either natural or anthropogenic disturbances at several timescales.     

生态系统文化服务研究进展——基于CiteSpace的可视化分析

生态系统文化服务（cultural ecosystem services, CES）作为一种重要的生态系统服务，是连接社会系统与自然系统的桥梁．以Web of Science平台的3 229条面板数据为样本，运用信息可视化软件CiteSpace进行了共被引分析、关键词共线分析、突现关键词检测等．结果表明:1）研究经历了萌芽期和发展期，其中萌芽期为1997—2005年，发展期又划分为缓慢发展期（2006—2012年）和快速发展期（2013—2019年），主要围绕概念框架、价值评估和制图、景观管理应用等方面开展；2）研究群体呈现大分散、小集中的特点，部分学者形成学术共同体，研究区主要分布在欧洲；3）价值评估一直是CES研究不变的主题，但在评估过程中，越来越注重利益相关者偏好和决策导向，与其他服务之间的权衡成为近年研究热点．      

产业集群生态系统协同演化的环分析

产业集群具有生态群落的特性,嵌入于企业生态系统之中,在集群企业种群内部企业间,集群企业种群间,以及产业集群与外部环境之间进行多层次的协同演化。各层次的协同演化促使产业集群随着环境的变化进行良性的演变和持续的发展,并获得较强的集群稳定性。还运用环分析方法对产业集群生态系统的稳定性进行判定。     

海南岛中部山区次生林生态特征研究

研究了海南岛中部山区典型的次生林群落特征和优势种枫香的种群结构特征．结果表明：海南岛中部山区的枫香种群所在的群落是演替过渡群落,在群落中枫香占有绝对优势,但随着群落的发展,枫香种群的发育表现出随着次生林年龄的增大而逐渐衰退的典型演替系列过渡种的生物学特性,最终会被群落中的后期演替种所代替．     

呼伦贝尔草原群落生产力与物种多样性关系分析

在呼伦贝草原选取5种代表性草原群落类型,采用取样调查的方法,探讨自然状况下草原植被群落生产力与物种多样性的关系,旨在为草原生态系统功能的恢复和草原有效管理提供理论依据.研究结果表明:大针茅+糙隐子草群落、羊草+克氏针茅群落、克氏针茅+羊草群落和贝加尔针茅+羊草群落多样性指数(物种丰富度指数、Simpson指数和Shannon-Winner指数)与生产力均呈显著正相关关系,但是羊草+寸草苔草甸草原物种多样性指数与生产力没有显著性相关关系.群落生产力受物种多样性、物种组成和环境因子等多因素共同影响,不仅仅是由物种多样性决定.     

Prolonged suppression of ecosystem carbon dioxide uptake after an anomalously warm year

Terrestrial ecosystems control carbon dioxide fluxes to and from the atmosphere through photosynthesis and respiration, a balance between net primary productivity and heterotrophic respiration, that determines whether an ecosystem is sequestering carbon or releasing it to the atmosphere. Global and site-specific data sets have demonstrated that climate and climate variability influence biogeochemical processes that determine net ecosystem carbon dioxide exchange (NEE) at multiple timescales. Experimental data necessary to quantify impacts of a single climate variable, such as temperature anomalies, on NEE and carbon sequestration of ecosystems at interannual timescales have been lacking. This derives from an inability of field studies to avoid the confounding effects of natural intra-annual and interannual variability in temperature and precipitation. Here we present results from a four-year study using replicate 12,000-kg intact tallgrass prairie monoliths located in four 184-m(3) enclosed lysimeters. We exposed 6 of 12 monoliths to an anomalously warm year in the second year of the study and continuously quantified rates of ecosystem processes, including NEE. We find that warming decreases NEE in both the extreme year and the following year by inducing drought that suppresses net primary productivity in the extreme year and by stimulating heterotrophic respiration of soil biota in the subsequent year. Our data indicate that two years are required for NEE in the previously warmed experimental ecosystems to recover to levels measured in the control ecosystems. This time lag caused net ecosystem carbon sequestration in previously warmed ecosystems to be decreased threefold over the study period, compared with control ecosystems. Our findings suggest that more frequent anomalously warm years, a possible consequence of increasing anthropogenic carbon dioxide levels, may lead to a sustained decrease in carbon dioxide uptake by terrestrial ecosystems.     

Stability of alpine meadow ecosystem on the Qinghai- Tibetan Plateau

THE QINGHAI-TIBETAN PLATEAU PLAYS AN IMPORTANT ROLE IN THE ATMOSPHERIC CIRCULATION AND REGIONAL MONSOON CLIMATE, WHICH HAS GREAT INFLUENCE ON THE REGIONAL AND GLOBAL CLIMATE. THUS, THE CHANGE OF THE QINGHAI-TIBETAN PLATEAU ECOSYSTEM IS EXPECTED TO AFFECT …     

甘肃兴隆山不同演替阶段典型森林群落的凋落物动态

【目的】分析不同演替阶段典型森林群落凋落物的量、组成特征及月动态,了解兴隆山森林生态系统碳贮量和养分循环状况。【方法】采用凋落物收集器法,对甘肃兴隆山森林演替阶段的3种典型森林群落针阔混交林(山杨(Populus davidiana)-白桦(Betula platyphylla)-青杄(Picea wilosonii)林)和暗针叶林(青杄-灌木林和青杄-箭竹(Fargesia nitida)-苔藓林)的凋落物量、组分、月动态进行了观测与研究。【结果】3种典型森林群落年凋落物量5 534.48～7 951.25 kg/hm²,大小排序为:山杨-白桦-青杄林>青杄-灌木林>青杄-箭竹-苔藓林,针阔混交林高于暗针叶林; 凋落量随森林正向演替的进行而不断减少。山杨-白桦-青杄林中以叶(44.91%)、杂物(20.53%)、枝(15.86%)、果(14.74%)为主,青杄-灌木林中以叶(41.22%)、杂物(23.58%)、枝(18.53%)、果(13.32%)为主,青杄-箭竹-苔藓林中以叶(37.48%)、杂物(27.51%)、枝(22.35%)为主; 在叶凋落物中,针阔混交林以阔叶为主,暗针叶林则以针叶为主。3种典型森林群落凋落量动态模式均为双峰型,但最高峰和最低峰出现时期有所不同,针阔混交林最高峰在10月,最低峰在7月; 暗针叶林最高峰在4—5月,最低峰在8—9月。针叶凋落量动态模式呈双峰型,高峰期出现在4月和10月; 阔叶、杂物、枝、果和花凋落动态模式呈单峰型,阔叶最高峰在10月,杂物、枝和果在4—5月,花在5—6月; 树皮凋落动态无明显变化规律。【结论】森林演替对凋落量及其凋落物组成影响明显; 随森林由阳性落叶阔叶林向阴性针叶林方向演替,森林年凋落量逐渐变小; 阔叶凋落量所占比例逐渐减小,而针叶所占比例逐渐增加。     

Diversity peaks at intermediate productivity in a laboratory microcosm

The species diversity of natural communities is often strongly related to their productivity. The pattern of this relationship seems to vary: diversity is known to increase monotonically with productivity, to decrease monotonically with productivity, and to be unimodally related to productivity, with maximum diversity occurring at intermediate levels of productivity. The mechanism underlying these patterns remains obscure, although many possibilities have been suggested. Here we outline a simple mechanism--involving selection in a heterogeneous environment--to explain these patterns, and test it using laboratory cultures of the bacterium Pseudomonas fluorescens. We grew diverse cultures over a wide range of nutrient concentrations, and found a strongly unimodal relationship between diversity and productivity in heterogeneous, but not in homogeneous, environments. Our result provides experimental evidence that the unimodal relationship often observed in natural communities can be caused by selection for specialized types in a heterogeneous environment.     

Climate change decreases aquatic ecosystem productivity of Lake Tanganyika,Africa

Although the effects of climate warming on the chemical and physical properties of lakes have been documented, biotic and ecosystem-scale responses to climate change have been only estimated or predicted by manipulations and models. Here we present evidence that climate warming is diminishing productivity in Lake Tanganyika, East Africa. This lake has historically supported a highly productive pelagic fishery that currently provides 25-40% of the animal protein supply for the populations of the surrounding countries. In parallel with regional warming patterns since the beginning of the twentieth century, a rise in surface-water temperature has increased the stability of the water column. A regional decrease in wind velocity has contributed to reduced mixing, decreasing deep-water nutrient upwelling and entrainment into surface waters. Carbon isotope records in sediment cores suggest that primary productivity may have decreased by about 20%, implying a roughly 30% decrease in fish yields. Our study provides evidence that the impact of regional effects of global climate change on aquatic ecosystem functions and services can be larger than that of local anthropogenic activity or overfishing.