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.     

Compartments revealed in food-web structure

Compartments in food webs are subgroups of taxa in which many strong interactions occur within the subgroups and few weak interactions occur between the subgroups. Theoretically, compartments increase the stability in networks, such as food webs. Compartments have been difficult to detect in empirical food webs because of incompatible approaches or insufficient methodological rigour. Here we show that a method for detecting compartments from the social networking science identified significant compartments in three of five complex, empirical food webs. Detection of compartments was influenced by food web resolution, such as interactions with weights. Because the method identifies compartmental boundaries in which interactions are concentrated, it is compatible with the definition of compartments. The method is rigorous because it maximizes an explicit function, identifies the number of non-overlapping compartments, assigns membership to compartments, and tests the statistical significance of the results. A graphical presentation reveals systemic relationships and taxa-specific positions as structured by compartments. From this graphic, we explore two scenarios of disturbance to develop a hypothesis for testing how compartmentalized interactions increase stability in food webs.     

Transient dynamics of an altered large marine ecosystem

Overfishing of large-bodied benthic fishes and their subsequent population collapses on the Scotian Shelf of Canada's east coast and elsewhere resulted in restructuring of entire food webs now dominated by planktivorous, forage fish species and macroinvertebrates. Despite the imposition of strict management measures in force since the early 1990s, the Scotian Shelf ecosystem has not reverted back to its former structure. Here we provide evidence of the transient nature of this ecosystem and its current return path towards benthic fish species domination. The prolonged duration of the altered food web, and its current recovery, was and is being governed by the oscillatory, runaway consumption dynamics of the forage fish complex. These erupting forage species, which reached biomass levels 900% greater than those prevalent during the pre-collapse years of large benthic predators, are now in decline, having outstripped their zooplankton food supply. This dampening, and the associated reduction in the intensity of predation, was accompanied by lagged increases in species abundances at both lower and higher trophic levels, first witnessed in zooplankton and then in large-bodied predators, all consistent with a return towards the earlier ecosystem structure. We conclude that the reversibility of perturbed ecosystems can occur and that this bodes well for other collapsed fisheries.     

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.     

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.     

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.     

Paleodietary changes by penguins and seals in association with Antarctic climate and sea ice extent

Positioned near the top of the food web, the dietary composition of Antarctic penguins and seals can be an excellent indicator of the regional food web and thus the status of the marine ecosystem. The dietary composition of modern penguins and seals has been well investigated; a long-term time series of data on penguin and seal diets, however, are rare. Such data, especially any predating the initiation of human harvesting of fish, whales and seals in Antarctica, are crucial for understanding and predicting responses of regional marine food webs to natural climate changes. Here we review recent progress on research of paleodietary change in Antarctic penguins and seals, specifically the Adelie penguin （Pygoscelis adeliae） and Antarctic fur seal （Arctocephalus gazella）. These studies indicate that the dietary changes of penguins correspond quite well with fluctuations in climate and sea ice extent during the Holocene. The depleted δ15N ratios found in modern Adelie penguins support the ＂krill surplus hypothesis＂ in relation to historic human depletion of krilleating fish, seals and whales.     

Overcompensation and population cycles in an ungulate.

Although theoretical studies show that overcompensatory density-dependent mechanisms can potentially generate regular or chaotic fluctuations in animal numbers, the majority of realistic single-species models of invertebrate populations are not overcompensatory enough to cause sustained population cycles. The possibility that overcompensation may generate cycles or chaos in vertebrate populations has seldom been considered. Here we show that highly overcompensatng density-dependent mortality can generate recurrent population crashes consistent with those observed in a naturally limited population of Soay sheep. The observed interval of three or more years between crashes points to sharp 'focusing' of mortality over a narrow range of population density.     

Symbiotic fungal endophytes control insect host-parasite interaction webs

Symbiotic microorganisms that live intimately associated with terrestrial plants affect both the quantity and quality of resources, and thus the energy supply to consumer populations at higher levels in the food chain. Empirical evidence on resource limitation of food webs points to primary productivity as a major determinant of consumer abundance and trophic structure. Prey quality plays a critical role in community regulation. Plants infected by endophytic fungi are known to be chemically protected against herbivore consumption. However, the influence of this microbe-plant association on multi-trophic interactions remains largely unexplored. Here we present the effects of fungal endophytes on insect food webs that reflect limited energy transfer to consumers as a result of low plant quality, rather than low productivity. Herbivore-parasite webs on endophyte-free grasses show enhanced insect abundance at alternate trophic levels, higher rates of parasitism, and increased dominance by a few trophic links. These results mirror predicted effects of increased productivity on food-web dynamics. Thus 'hidden' microbial symbionts can have community-wide impacts on the pattern and strength of resource-consumer interactions.     

Experimental demonstration of chaos in a microbial food web

Discovering why natural population densities change over time and vary with location is a central goal of ecological and evolutional disciplines. The recognition that even simple ecological systems can undergo chaotic behaviour has made chaos a topic of considerable interest among theoretical ecologists. However, there is still a lack of experimental evidence that chaotic behaviour occurs in the real world of coexisting populations in multi-species systems. Here we study the dynamics of a defined predator-prey system consisting of a bacterivorous ciliate and two bacterial prey species. The bacterial species preferred by the ciliate was the superior competitor. Experimental conditions were kept constant with continuous cultivation in a one-stage chemostat. We show that the dynamic behaviour of such a two-prey, one-predator system includes chaotic behaviour, as well as stable limit cycles and coexistence at equilibrium. Changes in the population dynamics were triggered by changes in the dilution rates of the chemostat. The observed dynamics were verified by estimating the corresponding Lyapunov exponents. Such a defined microbial food web offers a new possibility for the experimental study of deterministic chaos in real biological systems.     

High-amplitude fluctuations and alternative dynamical states of midges in Lake Myvatn

Complex dynamics are often shown by simple ecological models and have been clearly demonstrated in laboratory and natural systems. Yet many classes of theoretically possible dynamics are still poorly documented in nature. Here we study long-term time-series data of a midge, Tanytarsus gracilentus (Diptera: Chironomidae), in Lake Myvatn, Iceland. The midge undergoes density fluctuations of almost six orders of magnitude. Rather than regular cycles, however, these fluctuations have irregular periods of 4-7 years, indicating complex dynamics. We fit three consumer-resource models capable of qualitatively distinct dynamics to the data. Of these, the best-fitting model shows alternative dynamical states in the absence of environmental variability; depending on the initial midge densities, the model shows either fluctuations around a fixed point or high-amplitude cycles. This explains the observed complex population dynamics: high-amplitude but irregular fluctuations occur because stochastic variability causes the dynamics to switch between domains of attraction to the alternative states. In the model, the amplitude of fluctuations depends strongly on minute resource subsidies into the midge habitat. These resource subsidies may be sensitive to human-caused changes in the hydrology of the lake, with human impacts such as dredging leading to higher-amplitude fluctuations. Tanytarsus gracilentus is a key component of the Myvatn ecosystem, representing two-thirds of the secondary productivity of the lake and providing vital food resources to fish and to breeding bird populations. Therefore the high-amplitude, irregular fluctuations in midge densities generated by alternative dynamical states dominate much of the ecology of the lake.     

一混沌模型的系综模拟研究

用随机模拟方法研究了一化学混沌模型的介观动力学 ,对该混沌模型的系综模拟发现 ,在这种不稳定运动中存在强烈的内部涨落 ,然而由于混沌运动整体上的稳定性 ,使得系综中的代表点被限制在混沌吸引子上 ,并且单个代表点形成的随机轨道很好地保持了确定性混沌吸引子的基本特征     

Universal scaling relations in food webs

The structure of ecological communities is usually represented by food webs. In these webs, we describe species by means of vertices connected by links representing the predations. We can therefore study different webs by considering the shape (topology) of these networks. Comparing food webs by searching for regularities is of fundamental importance, because universal patterns would reveal common principles underlying the organization of different ecosystems. However, features observed in small food webs are different from those found in large ones. Furthermore, food webs (except in isolated cases) do not share general features with other types of network (including the Internet, the World Wide Web and biological webs). These features are a small-world character and a scale-free (power-law) distribution of the degree (the number of links per vertex). Here we propose to describe food webs as transportation networks by extending to them the concept of allometric scaling (how branching properties change with network size). We then decompose food webs in spanning trees and loop-forming links. We show that, whereas the number of loops varies significantly across real webs, spanning trees are characterized by universal scaling relations.     

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.     

Stability criteria for complex ecosystems

Forty years ago, May proved that sufficiently large or complex ecological networks have a probability of persisting that is close to zero, contrary to previous expectations. May analysed large networks in which species interact at random. However, in natural systems pairs of species have well-defined interactions (for example predator-prey, mutualistic or competitive). Here we extend May's results to these relationships and find remarkable differences between predator-prey interactions, which are stabilizing, and mutualistic and competitive interactions, which are destabilizing. We provide analytic stability criteria for all cases. We use the criteria to prove that, counterintuitively, the probability of stability for predator-prey networks decreases when a realistic food web structure is imposed or if there is a large preponderance of weak interactions. Similarly, stability is negatively affected by nestedness in bipartite mutualistic networks. These results are found by separating the contribution of network structure and interaction strengths to stability. Stable predator-prey networks can be arbitrarily large and complex, provided that predator-prey pairs are tightly coupled. The stability criteria are widely applicable, because they hold for any system of differential equations.     

Evolution of species interactions in a biofilm community

Biofilms are spatially structured communities of microbes whose function is dependent on a complex web of symbiotic interactions. Localized interactions within these assemblages are predicted to affect the coexistence of the component species, community structure and function, but there have been few explicit empirical analyses of the evolution of interactions. Here we show, with the use of a two-species community, that selection in a spatially structured environment leads to the evolution of an exploitative interaction. Simple mutations in the genome of one species caused it to adapt to the presence of the other, forming an intimate and specialized association. The derived community was more stable and more productive than the ancestral community. Our results show that evolution in a spatially structured environment can stabilize interactions between species, provoke marked changes in their symbiotic nature and affect community function.     

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.     

土壤食物网调控及其生态功能研究的困境与思考

土壤食物网精准调控是探究和发挥土壤生物生态功能的关键和难点.综述了已有的土壤食物网直接和间接调控方法,讨论了土壤食物网生态功能研究现状及面临的困境,如常用的直接调控措施很难在野外大规模应用;间接调控土壤食物网内在机制复杂,调控效果不易把控;土壤食物网全图模糊不清,关键类群及其权重难确定;土壤食物网的物质循环与能流难量化...     

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.