Ecosystem-level patterns of primary productivity and herbivory in terrestrial habitats

Ecosystems are structurally organized as food webs within which energy is transmitted between trophic levels and dissipated into the environment. Energy flow between two trophic levels is given by the amount of production at the lower level and by the proportion of production that is consumed, assimilated and respired at the higher level. Considerable evidence indicates that food-web structure varies predictably in different habitats, but much less is known about quantitative relationships among food web fluxes. Many of the energetic properties of herbivores in African game parks are associated with rainfall and, by inference, with net primary productivity. Respiratory costs per unit production at the consumer trophic level are higher for homeotherms than for heterotherms. Plant secondary chemicals affect herbivore dietary choices and the allocation of plant resources to those chemicals varies with resource availability. How these phenomena are translated into ecosystem fluxes is unknown. We present evidence that herbivore biomass, consumption and productivity are closely correlated with plant productivity, suggesting that the latter is a principal integrator and indicator of functional processes in food webs.     

Unsaturated fatty acid content in seston and tropho-dynamic coupling in lakes

Determining the factors that control food web interactions is a key issue in ecology. The empirical relationship between nutrient loading (total phosphorus) and phytoplankton standing stock (chlorophyll a) in lakes was described about 30 years ago and is central for managing surface water quality. The efficiency with which biomass and energy are transferred through the food web and sustain the production of higher trophic levels (such as fish) declines with nutrient loading and system productivity, but the underlying mechanisms are poorly understood. Here we show that in seston (fine particles in water) during summer, specific omega3-polyunsaturated fatty acids (omega3-PUFAs), which are important for zooplankton, are significantly correlated to the trophic status of the lake. The omega3-PUFAs octadecatetraenoic acid, eicosapentaenoic acid (EPA) and docosahexaenoic acid, but not alpha-linolenic acid, decrease on a double-logarithmic scale with increasing total phosphorus. By combining the empirical relationship between EPA-to-carbon content and total phosphorus with functional models relating EPA-to-carbon content to the growth and egg production of daphnids, we predict secondary production for this key consumer. Thus, the decreasing efficiency in energy transfer with increasing lake productivity can be explained by differences in omega3-PUFA-associated food quality at the plant-animal interface.     

Ecosystem consequences of species richness and composition in pond food webs

Resolving current concerns about the role of biodiversity on ecosystems calls for understanding the separate roles of changes in species numbers and of composition. Recent work shows that primary productivity often, but not always, saturates with species richness within single trophic levels. However, any interpretation of such patterns must consider that variation in biodiversity is necessarily associated with changes in species composition (identity), and that changes in biodiversity often occur across multiple trophic levels. Here we present results from a mesocosm experiment in which we independently manipulated species richness and species composition across multiple trophic levels in pond food webs. In contrast to previous studies that focused on single trophic levels, we found that productivity is either idiosyncratic or increases with respect to species richness, and that richness influences trophic structure. However, the composition of species within richness levels can have equally or more marked effects on ecosystems than average effects of richness per se. Indirect evidence suggests that richness and associated changes in species composition affect ecosystem attributes through indirect effects and trophic interactions among species, features that are highly characteristic of natural, complex ecosystems.     

Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment

Biodiversity is rapidly declining, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above- and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades.     

Trophic cascades across ecosystems

Predation can be intense, creating strong direct and indirect effects throughout food webs. In addition, ecologists increasingly recognize that fluxes of organisms across ecosystem boundaries can have major consequences for community dynamics. Species with complex life histories often shift habitats during their life cycles and provide potent conduits coupling ecosystems. Thus, local interactions that affect predator abundance in one ecosystem (for example a larval habitat) may have reverberating effects in another (for example an adult habitat). Here we show that fish indirectly facilitate terrestrial plant reproduction through cascading trophic interactions across ecosystem boundaries. Fish reduce larval dragonfly abundances in ponds, leading to fewer adult dragonflies nearby. Adult dragonflies consume insect pollinators and alter their foraging behaviour. As a result, plants near ponds with fish receive more pollinator visits and are less pollen limited than plants near fish-free ponds. Our results confirm that strong species interactions can reverberate across ecosystems, and emphasize the importance of landscape-level processes in driving local species interactions.     

Dimensionality of consumer search space drives trophic interaction strengths

Trophic interactions govern biomass fluxes in ecosystems, and stability in food webs. Knowledge of how trophic interaction strengths are affected by differences among habitats is crucial for understanding variation in ecological systems. Here we show how substantial variation in consumption-rate data, and hence trophic interaction strengths, arises because consumers tend to encounter resources more frequently in three dimensions (3D) (for example, arboreal and pelagic zones) than two dimensions (2D) (for example, terrestrial and benthic zones). By combining new theory with extensive data (376 species, with body masses ranging from 5.24 × 10(-14) kg to 800 kg), we find that consumption rates scale sublinearly with consumer body mass (exponent of approximately 0.85) for 2D interactions, but superlinearly (exponent of approximately 1.06) for 3D interactions. These results contradict the currently widespread assumption of a single exponent (of approximately 0.75) in consumer-resource and food-web research. Further analysis of 2,929 consumer-resource interactions shows that dimensionality of consumer search space is probably a major driver of species coexistence, and the stability and abundance of populations.     

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.     

瓜类蔬菜昆虫群落（包括蛛形纲和软体动物）的研究 Ⅰ．群落的组成及结构

本文是重庆市郊瓜类蔬菜昆虫群落研究结果的第一部分,主要报道瓜类昆虫落（包括蛛形纲和软体动物门中的一些有害生物及天敌,下同）的组成和结构,初步查明对瓜类蔬菜有直接或间接危害的动物有53种,分属于3纲,14目中的32个科;天敌38种,涉及2纲,9目中的21个科,另外还有传粉昆虫和杂食性昆虫各1种,分属于2个科,在初步弄清群落内物种间营养关系的基础上,用营养物种的概念探讨了不同营养物种数,不同类型链节数,链节总数与物种数之间的关系,表明瓜类昆虫群落属于发育不成熟,结构较为简单的群落,本文还同时应用基于食物网映射矩阵的匹配系数和基于物种及其多度的群落系数对5种瓜类昆虫群落相似性进行了比较。     

Structural asymmetry and the stability of diverse food webs

Untangling the influence of human activities on food-web stability and persistence is complex given the large numbers of species and overwhelming number of interactions within ecosystems. Although biodiversity has been associated with stability, the actual structures and processes that confer stability to diverse food webs remain largely unknown. Here we show that real food webs are structured such that top predators act as couplers of distinct energy channels that differ in both productivity and turnover rate. Our theoretical analysis shows that coupled fast and slow channels convey both local and non-local stability to food webs. Alarmingly, the same human actions that have been implicated in the loss of biodiversity also directly erode the very structures and processes that we show to confer stability on food webs.     

Predator diversity dampens trophic cascades

Food web complexity is thought to weaken the strength of terrestrial trophic cascades in which strong impacts of natural enemies on herbivores cascade to influence primary production indirectly. Predator diversity can enhance food web complexity because predators may feed on each other and on shared prey. In such cases, theory suggests that the impact of predation on herbivores relaxes and cascading effects on basal resources are dampened. Despite this view, no empirical studies have explicitly investigated the role of predator diversity in mediating primary productivity in a natural terrestrial system. Here we compare, in a coastal marsh community, impacts of arthropod predators on herbivores and plant productivity between a simple food web with a single predator species and a complex food web with a diverse predator assemblage. We show that enhancing predator diversity dampens enemy effects on herbivores and weakens trophic cascades. Consequently, changes in diversity at higher trophic levels can significantly alter ecosystem function in natural systems.     

Simple rules yield complex food webs

Several of the most ambitious theories in ecology describe food webs that document the structure of strong and weak trophic links that is responsible for ecological dynamics among diverse assemblages of species. Early mechanism-based theory asserted that food webs have little omnivory and several properties that are independent of species richness. This theory was overturned by empirical studies that found food webs to be much more complex, but these studies did not provide mechanistic explanations for the complexity. Here we show that a remarkably simple model fills this scientific void by successfully predicting key structural properties of the most complex and comprehensive food webs in the primary literature. These properties include the fractions of species at top, intermediate and basal trophic levels, the means and variabilities of generality, vulnerability and food-chain length, and the degrees of cannibalism, omnivory, looping and trophic similarity. Using only two empirical parameters, species number and connectance, our 'niche model' extends the existing 'cascade model and improves its fit ten-fold by constraining species to consume a contiguous sequence of prey in a one-dimensional trophic niche.     

高温胁迫对植食性昆虫影响研究进展

昆虫作为变温动物类群,极易遭受高温胁迫的影响。研究高温胁迫对植食性昆虫的影响,可为气候变暖背景下农林害虫的种群动态监测及地理适生区预测提供重要理论依据。笔者从植食性昆虫个体、种间互作关系、种群及群落3个层面,综述了高温胁迫对植食性昆虫影响的国内外进展。研究发现,高温胁迫会对植食性昆虫个体的生长发育、繁殖及生理生化带来负面效应,但昆虫自身也会进化出一系列基于形态、行为及基因表达的热胁迫响应机制。高温胁迫对寄主植物、天敌、共生微生物的影响也能够通过种间互作传递给植食性昆虫,并导致寄主植物-植食性昆虫-天敌3级营养关系发生变化。同时,高温胁迫还能通过影响昆虫个体及种间关系,间接影响昆虫的种群动态、种群多样性及其生态功能,并可能导致植食性昆虫的种群大爆发或衰退。最后,笔者认为此领域未来研究方向为:在个体层面上,应优化高温胁迫处理方式并综合考虑干旱和降水等环境因子,开展农林害虫种群的长期野外监测,应注重高温胁迫对昆虫参与生长发育繁殖功能基因的影响;在种间关系层面上,应关注高温胁迫对昆虫关联的复杂多样的食物网及互作体系的影响;此外,应结合高温胁迫对不同昆虫种类自身生理特性和行为模式的影响,阐明温度升高对昆虫群落的影响。     

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.     

Altered performance of forest pests under atmospheres enriched by CO2 and O3

Human activity causes increasing background concentrations of the greenhouse gases CO2 and O3. Increased levels of CO2 can be found in all terrestrial ecosystems. Damaging O3 concentrations currently occur over 29% of the world's temperate and subpolar forests but are predicted to affect fully 60% by 2100 (ref. 3). Although individual effects of CO2 and O3 on vegetation have been widely investigated, very little is known about their interaction, and long-term studies on mature trees and higher trophic levels are extremely rare. Here we present evidence from the most widely distributed North American tree species, Populus tremuloides, showing that CO2 and O3, singly and in combination, affected productivity, physical and chemical leaf defences and, because of changes in plant quality, insect and disease populations. Our data show that feedbacks to plant growth from changes induced by CO2 and O3 in plant quality and pest performance are likely. Assessments of global change effects on forest ecosystems must therefore consider the interacting effects of CO2 and O3 on plant performance, as well as the implications of increased pest activity.     

Single-species models for many-species food webs

Most species live in species-rich food webs; yet, for a century, most mathematical models for population dynamics have included only one or two species. We ask whether such models are relevant to the real world. Two-species population models of an interacting consumer and resource collapse to one-species dynamics when recruitment to the resource population is unrelated to resource abundance, thereby weakening the coupling between consumer and resource. We predict that, in nature, generalist consumers that feed on many species should similarly show one-species dynamics. We test this prediction using cyclic populations, in which it is easier to infer underlying mechanisms, and which are widespread in nature. Here we show that one-species cycles can be distinguished from consumer resource cycles by their periods. We then analyse a large number of time series from cyclic populations in nature and show that almost all cycling, generalist consumers examined have periods that are consistent with one-species dynamics. Thus generalist consumers indeed behave as if they were one-species populations, and a one-species model is a valid representation for generalist population dynamics in many-species food webs.     

The trophic fingerprint of marine fisheries

Biodiversity indicators provide a vital window on the state of the planet, guiding policy development and management. The most widely adopted marine indicator is mean trophic level (MTL) from catches, intended to detect shifts from high-trophic-level predators to low-trophic-level invertebrates and plankton-feeders. This indicator underpins reported trends in human impacts, declining when predators collapse ("fishing down marine food webs") and when low-trophic-level fisheries expand ("fishing through marine food webs"). The assumption is that catch MTL measures changes in ecosystem MTL and biodiversity. Here we combine model predictions with global assessments of MTL from catches, trawl surveys and fisheries stock assessments and find that catch MTL does not reliably predict changes in marine ecosystems. Instead, catch MTL trends often diverge from ecosystem MTL trends obtained from surveys and assessments. In contrast to previous findings of rapid declines in catch MTL, we observe recent increases in catch, survey and assessment MTL. However, catches from most trophic levels are rising, which can intensify fishery collapses even when MTL trends are stable or increasing. To detect fishing impacts on marine biodiversity, we recommend greater efforts to measure true abundance trends for marine species, especially those most vulnerable to fishing.     

Allometric degree distributions facilitate food-web stability

In natural ecosystems, species are linked by feeding interactions that determine energy fluxes and create complex food webs. The stability of these food webs enables many species to coexist and to form diverse ecosystems. Recent theory finds predator-prey body-mass ratios to be critically important for food-web stability. However, the mechanisms responsible for this stability are unclear. Here we use a bioenergetic consumer-resource model to explore how and why only particular predator-prey body-mass ratios promote stability in tri-trophic (three-species) food chains. We find that this 'persistence domain' of ratios is constrained by bottom-up energy availability when predators are much smaller than their prey and by enrichment-driven dynamics when predators are much larger. We also find that 97% of the tri-trophic food chains across five natural food webs exhibit body-mass ratios within the predicted persistence domain. Further analyses of randomly rewired food webs show that body mass and allometric degree distributions in natural food webs mediate this consistency. The allometric degree distributions hold that the diversity of species' predators and prey decreases and increases, respectively, with increasing species' body masses. Our results demonstrate how simple relationships between species' body masses and feeding interactions may promote the stability of complex food webs.     

Variance in ecological consumer-resource interactions

Food-web models use the effect size of trophic interactions to predict consumer-resource dynamics. These models anticipate that strong effects of consumers increase spatial and temporal variability in abundance of species, whereas weak effects dampen fluctuations. Empirical evidence indicates that opposite patterns may occur in natural assemblages. Here I show that spatial variance in the distribution of resource populations is sensitive to changes in the variance of the trophic interaction, in addition to the mean effect of consumers, relative to other causes of spatial variability. Simulations indicate that both strong and weak direct effects of consumers can promote spatial variability in abundance of resources, but only trophic interactions with a large mean effect size can reduce variation. Predictions of the model agree with the results of repeated field experiments and are consistent with data from published consumer-resource interactions, proving to be robust across widely varying environmental conditions and species' life histories. Thus, food-web models that embody variance in trophic interactions may have increased capacity to explain the wide range of effects of consumers documented in empirical studies.     

Unified spatial scaling of species and their trophic interactions

Two largely independent bodies of scaling theory address the quantitative relationships between habitat area, species diversity and trophic interactions. Spatial theory within macroecology addresses how species richness scales with area in landscapes, while typically ignoring interspecific interactions. Complexity theory within community ecology addresses how trophic links scale with species richness in food webs, while typically ignoring spatial considerations. Recent studies suggest unifying these theories by demonstrating how spatial patterns influence food-web structure and vice versa. Here, we follow this suggestion by developing and empirically testing a more unified scaling theory. On the basis of power-law species-area relationships, we develop link-area and non-power-law link-species models that accurately predict how trophic links scale with area and species richness of microcosms, lakes and streams from community to metacommunity levels. In contrast to previous models that assume that species richness alone determines the number of trophic links, these models include the species' spatial distribution, and hence extend the domain of complexity theory to metacommunity scales. This generality and predictive success shows how complexity theory and spatial theory can be unified into a much more general theory addressing new domains of ecology.     

生态系统的可靠性及其稳定性的维持

 生态系统的营养结构是由营养级组成的.可靠性模型实验表明,营养级的可靠性取决于物种的失效率大小,物种丢失对物种丰富、失效率高的营养级的可靠性影响甚微,而对物种贫乏、失效率低的营养级略大,换言之,营养级的可靠性与物种的丰富度无紧密关系.其次,在营养级可靠性相同的条件下,简单系统的可靠性普遍略高于营养级多或食物链长的复杂生态系统.生物多样性与生态系统的稳定性没有必然联系,只要营养级还存在(无论剩下几个物种),那么按串并联方式组合成的生态系统就不会崩溃.生态系统的稳定性是冗余结构的稳定性.生态系统经常遭受干扰,但物种和个体的丢失可以通过来自其内外源源不断的冗余补充,以维持生态系统营养结构的稳定性.