Weighting and indirect effects identify keystone species in food webs

Species extinctions are accelerating globally, yet the mechanisms that maintain local biodiversity remain poorly understood. The extinction of species that feed on or are fed on by many others (i.e. ‘hubs’) has traditionally been thought to cause the greatest threat of further biodiversity loss. Very little attention has been paid to the strength of those feeding links (i.e. link weight) and the prevalence of indirect interactions. Here, we used a dynamical model based on empirical energy budget data to assess changes in ecosystem stability after simulating the loss of species according to various extinction scenarios. Link weight and/or indirect effects had stronger effects on food‐web stability than the simple removal of ‘hubs’, demonstrating that both quantitative fluxes and species dissipating their effects across many links should be of great concern in biodiversity conservation, and the potential for ‘hubs’ to act as keystone species may have been exaggerated to date.     

Using community viability analysis to identify fragile systems and keystone species

Owing to interdependences among species in ecological communities, the loss of one species can trigger a cascade of secondary extinctions with potentially dramatic effects on the functioning and stability of the community. It is, therefore, important to assess the risk and likely extent of secondary extinctions. Community viability analysis is a new technique that can be used to accomplish this goal. The analysis can also be used to identify fragile community structures and keystone species and, hence, to provide guidelines for conservation priorities. Here, we describe the principles underlying community viability analysis and review its contributions to our understanding of the response of ecological communities to species loss.     

Topological keystone species: measures of positional importance in food webs

The local extinction or large fluctuation in abundance of a species may seriously affect other species in the community. The effects spread through the community by direct and indirect interactions. The network perspective on ecology can help map the pathways of these effects, for food webs, the pathways of indirect trophic interactions. Indirect interactions typically decay in intensity as they spread. Therefore, there is a conceptual maximum range in topological space beyond which interactions have no effects, even though all species remain connected. Neither the local characteristics of species, nor the global characteristics of entire webs, suitably quantify this range. We therefore apply intermediate scale indices that reflect the limitations imposed by effect damping in networks. We present a complex analysis of the topological positional importance of species in the Chesapeake Bay web. This web is a carbon-flow network that represents trophic interactions. We present several different indices reflecting different properties and discuss which questions the different indices best answer. We look for the best indices for identifying the key players in ecosystem functioning. Our study contributes to the quantification of relative species importance and provides an exact and a priori determination of a class of candidate keystone species that can inform applied and conservation ecology as well as theoretical concerns.     

Potential oscillators and keystone modules in food webs

Food web theory suggests that the placement of a weak interaction is critical such that under some conditions even one well‐placed weak interaction can stabilise multiple strong interactions. This theory suggests that complex stable webs may be built from pivotal weak interactions such that the removal of even one to a few keystone interactions can have significant cascading impacts on whole system diversity and structure. However, the connection between weak interactions, derived from the theory of modular food web components, and keystone species, derived from empirical results, is not yet well understood. Here, we develop numerical techniques to detect potential oscillators hidden in complex food webs, and show that, both in random and real food webs, keystone consumer–resource interactions often operate to stabilise them. Alarmingly, this result suggests that nature frequently may be dangerously close to precipitous change with even the loss of one or a few weakly interacting species.     

Effects of environmental warming and drought on size‐structured soil food webs

Soil systems maintain important ecosystem processes crucial for plant life and food production. Especially agricultural systems are strongly affected by climate change due to low vegetation cover associated with high temperatures and drought. Nevertheless, the response of soil systems to climate change is little explored. We used microcosms with a simplified soil community to address effects of climate change using independent temperature and dryness gradients and addressed their effects on top–down control and litter decomposition. The community consisted of maize litter as a basal resource, fungi, springtails and as top predators mites and centipedes. As the body‐size structure is of high importance for communities, we included differently‐sized springtails and predator species. After seven weeks, the experiment was terminated, and the impact of climate change on direct feeding interactions and indirect effects across trophic levels was analysed. With increasing temperature and dryness, consumption rates increased, thereby amplifying the negative influence of consumer populations on their resources. Hence, these climate‐change variables increased the top–down control of 1) predators (mainly mites) on springtails and 2) fungi on litter decomposition. In addition, we found that the climate‐change variables strengthened trophic cascades from predators on fungi whose density was thus increasingly decoupled from top–down control by their springtail consumers. Their increased decomposition rates are of high importance for carbon cycling and may result in accelerated nutrient turnover. In conclusion, our results suggest that climate change may strongly influence the structure and functioning of soil systems by strengthening consumption rates and trophic cascades, which will have far reaching consequences for the nutrient turnover and productivity of agricultural ecosystems.     

Differential responses of size‐based functional groups to bottom–up and top–down perturbations in pelagic food webs: a meta‐analysis

We performed a meta‐analysis of 31 lake mesocosm experiments to investigate differences in the responses of pelagic food chains and food webs to nutrient enrichment and fish presence. Trophic levels were divided into size‐based functional groups (phytoplankton into highly edible and poorly edible algae, and zooplankton into small herbivores, large herbivores and omnivorous zooplankton) in the food webs. Our meta‐analysis shows that 1) nutrient enrichment has a positive effect on phytoplankton and zooplankton, while fish presence has a positive effect on phytoplankton and a negative effect on zooplankton in the food chains; 2) nutrient enrichment has a positive effect on highly edible algae and small herbivores, but no effect on poorly edible algae, large herbivores and omnivorous zooplankton in the food webs. Planktivorous fish have a positive effect on highly edible algae and small herbivores, a negative effect on large herbivores and omnivorous zooplankton, and no effect on poorly edible algae. Our meta‐analysis confirms that nutrient enrichment and planktivorous fish affect functional groups differentially within trophic levels, revealing important changes in the functioning of food webs. The analysis of fish effects shows the well‐described trophic cascade in the food chain and reveals two trophic cascades in the food web: one transmitted by large herbivores that benefit highly edible phytoplankton, and one transmitted by omnivorous zooplankton that benefit small herbivores. Comparison between the responses of food webs and simple food chains also shows consistent biomass compensation between functional groups within trophic levels.     

Functional links and robustness in food webs

The robustness of ecosystems to species losses is a central question in ecology, given the current pace of extinctions and the many species threatened by human impacts, including habitat destruction and climate change. Robustness from the perspective of secondary extinctions has been addressed in the context of food webs to consider the complex network of species interactions that underlie responses to perturbations. In-silico removal experiments have examined the structural properties of food webs that enhance or hamper the robustness of ecosystems to species losses, with a focus on the role of hubs, the most connected species. Here we take a different approach and focus on the role of the connections themselves. We show that trophic links can be divided into functional and redundant based on their contribution to robustness. The analysis of empirical webs shows that hubs are not necessarily the most important species as they may hold many redundant links. Furthermore, the fraction of functional connections is high and constant across systems regardless of size and interconnectedness. The main consequence of this scaling pattern is that ecosystem robustness can be considerably reduced by species extinctions even when these do not result in any secondary extinctions. This introduces the possibility of tipping points in the collapse of ecosystems.     

Characterization of topological keystone species: Local,global and “meso-scale” centralities in food webs

An important question in the network representation of ecological systems is to determine how direct and indirect interactions between species determine the positional importance of species in the ecosystem. Here we present a quantitative analysis of the similarities and differences of six different topological centrality measures as indicators of keystone species in 17 food webs. These indicators account for local, global and “meso-scale” – intermediate between local and global – topological information about species in the food webs. Using factor analysis we shown that most of these centrality indices share a great deal of topological information, which range from 75% to 96%. A generalized keystone indicator is then proposed by considering the factor loadings of the six-centrality measures, which contains most of the information encoded by these indices. However, the individual ordering of species according to these criteria display significant differences in most food webs. We simulate the effects of species extinction by removing species ranked according to a local and a “meso-scale” centrality indicator. The differences observed on three network characteristics – size, average distance and clustering coefficient of the largest component – after the removal of the most central nodes indicate that the consideration of these indices have different impacts for the ranking of species with conservational biology purposes. The “meso-scale” indicator appears to play an important role in determining the relative importance of species in epidemic spread and parasitism rates.     

Keystone species and food webs

Different species are of different importance in maintaining ecosystem functions in natural communities. Quantitative approaches are needed to identify unusually important or influential, ‘keystone’ species particularly for conservation purposes. Since the importance of some species may largely be the consequence of their rich interaction structure, one possible quantitative approach to identify the most influential species is to study their position in the network of interspecific interactions. In this paper, I discuss the role of network analysis (and centrality indices in particular) in this process and present a new and simple approach to characterizing the interaction structures of each species in a complex network. Understanding the linkage between structure and dynamics is a condition to test the results of topological studies, I briefly overview our current knowledge on this issue. The study of key nodes in networks has become an increasingly general interest in several disciplines: I will discuss some parallels. Finally, I will argue that conservation biology needs to devote more attention to identify and conserve keystone species and relatively less attention to rarity.     

Parasites in food webs: the ultimate missing links

Parasitism is the most common consumer strategy among organisms, yet only recently has there been a call for the inclusion of infectious disease agents in food webs. The value of this effort hinges on whether parasites affect food‐web properties. Increasing evidence suggests that parasites have the potential to uniquely alter food‐web topology in terms of chain length, connectance and robustness. In addition, parasites might affect food‐web stability, interaction strength and energy flow. Food‐web structure also affects infectious disease dynamics because parasites depend on the ecological networks in which they live. Empirically, incorporating parasites into food webs is straightforward. We may start with existing food webs and add parasites as nodes, or we may try to build food webs around systems for which we already have a good understanding of infectious processes. In the future, perhaps researchers will add parasites while they construct food webs. Less clear is how food‐web theory can accommodate parasites. This is a deep and central problem in theoretical biology and applied mathematics. For instance, is representing parasites with complex life cycles as a single node equivalent to representing other species with ontogenetic niche shifts as a single node? Can parasitism fit into fundamental frameworks such as the niche model? Can we integrate infectious disease models into the emerging field of dynamic food‐web modelling? Future progress will benefit from interdisciplinary collaborations between ecologists and infectious disease biologists.     

Adaptive omnivory and species coexistence in tri-trophic food webs

The commonness of omnivory in natural communities is puzzling, because simple dynamic models of tri-trophic systems with omnivory are prone to species extinction. In particular, the intermediate consumer is frequently excluded by the omnivore at high levels of enrichment. It has been suggested that adaptive foraging by the omnivore may facilitate coexistence, because the intermediate consumer should persist more easily if it is occasionally dropped from the omnivore's diet. We explore theoretically how species permanence in tri-trophic systems is affected if the omnivore forages adaptively according to the "diet rule", i.e., feeds on the less profitable of its two prey species only if the more profitable one is sufficiently rare. We show that, compared to systems where omnivory is fixed, adaptive omnivory may indeed facilitate 3-species persistence. Counter to intuition, however, facilitation of 3-species coexistence requires that the intermediate consumer is a more profitable prey than the basal resource. Consequently, adaptive omnivory does not facilitate persistence of the intermediate consumer but enlarges the persistence region of the omnivore towards parameter space where a fixed omnivore would be excluded by the intermediate consumer. Overall, the positive effect of adaptive omnivory on 3-species persistence is, however, small. Generally, whether omnivory is fixed or adaptive, 3-species permanence is most likely when profitability (=conversion efficiency into omnivores) is low for basal resources and high for intermediate consumers.     

基于SURF指数识别海州湾食物网的关键饵料生物

关键种(keystone species)在生态系统中发挥着不可替代的重要作用,对于群落结构的稳定与演替起着决定性的作用。基于2011年3—12月在海州湾及其邻近海域进行的渔业资源底拖网调查资料以及胃含物分析数据和参考历史文献数据,以物种间的摄食关系作为基础,采用SURF(Supportive Role to Fishery ecosystems)指数识别海州湾食物网中的关键饵料生物。在矩阵中分析每个物种作为饵料生物为捕食者提供的摄食比例及捕食者数量,计算每个物种的关键指标(SURFi)的值,结果表明前五位数值较高的物种是细鳌虾(Leptochela gracilis)、毛虾(Acetessp.)、疣背宽额虾(Latreutes planirostris)、日本鼓虾(Alpheus japonicus)和鳀鱼(Engraulis japonicus),它们是海州湾的关键饵料生物,在整个食物网中起到关键的控制作用,它们的数量波动会对海州湾其他物种产生直接或间接的影响。加强关键饵料生物的保护,对于维持海州湾生态系统的健康和稳定至关重要。     

High‐resolution food webs based on nitrogen isotopic composition of amino acids

Food webs are known to have myriad trophic links between resource and consumer species. While herbivores have well‐understood trophic tendencies, the difficulties associated with characterizing the trophic positions of higher‐order consumers have remained a major problem in food web ecology. To better understand trophic linkages in food webs, analysis of the stable nitrogen isotopic composition of amino acids has been introduced as a potential means of providing accurate trophic position estimates. In the present study, we employ this method to estimate the trophic positions of 200 free‐roaming organisms, representing 39 species in coastal marine (a stony shore) and 38 species in terrestrial (a fruit farm) environments. Based on the trophic positions from the isotopic composition of amino acids, we are able to resolve the trophic structure of these complex food webs. Our approach reveals a high degree of trophic omnivory (i.e., noninteger trophic positions) among carnivorous species such as marine fish and terrestrial hornets.This information not only clarifies the trophic tendencies of species within their respective communities, but also suggests that trophic omnivory may be common in these webs.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Sex‐biased resource allocation in ovo in a sexually size‐dimorphic species

Sex‐biased resource allocation in eggs is increasingly recognized as one strategy oviparous mothers can employ to invest differentially in one sex, depending on nutritional requirements. Previous studies have used egg size as an index of nutrient allocation, but few have examined egg contents directly. We used molecular sexing of early‐stage ring‐billed gull Larus delawarensis embryos, a species with sexual size dimorphism, to test whether sex‐specific nutrient allocation occurs in ovo. Despite no sex difference in size, eggs with male embryos contained more albumen, while eggs with female embryos contained more yolk, lipid and non‐lipid (protein and carbohydrate). It is unclear why such sex‐biased resource allocation in ovo is utilized by ring‐billed gulls. However, our data indicate that a cursive examination of egg mass or size may not necessarily reflect nutrient allocation strategies mothers use in ovo, and that sex‐biased investment in ovo may be more widespread than currently appreciated.     

Using DNA‐based techniques to identify hybrids among linear‐leaved Potamogeton plants collected in China

Abstract It is well known that interspecific hybrids occur in the genus Potamogeton. The linear‐leaved Potamogeton species commonly have highly variable morphological characteristics. Their hybrids often show similar vegetative characters to their parental species and their identification based solely on morphology is not always conclusive. In order to clarify whether there are any hybrids from the linear‐leaved Potamogeton plants collected in China, we used internal transcribed spacers (ITS) of nuclear ribosomal DNA and chloroplast rbcL gene sequences to identify the hybrids. Using ITS sequence additivity, we identified four hybrids, namely P. orientalis (P. pusillus×P. oxyphyllus), P. pusillus×P. berchtoldii, P. foliosus×P. octandrus, and P. cristatus×P. octandrus. The latter three hybrids should be considered as new hybrids in Potamogeton. The maternal parents of the four hybrids were confirmed using chloroplast rbcL gene sequences.     

Reverse latitudinal trends in species richness of pitcher-plant food webs

Latitudinal patterns in species richness have been well documented for guilds and individual trophic groups, but comparable patterns for entire, multitrophic communities have not been described. We studied the entire food web that inhabits the water‐filled leaves of the pitcher plant Sarracenia purpurea across North America at two spatial scales: among sites and among leaves within sites. Contrary to the expectation, total species richness at both scales increased with latitude, because of increasing species richness at the lower trophic levels. This latitudinal pattern may be driven by a top‐down effect. The abundance of the mosquito Wyeomyia smithii, a ubiquitous top predator in this system, decreases from south to north and may permit greater species richness of prey trophic levels at higher latitudes.     

Connectance indicates the robustness of food webs when subjected to species loss

Many ecologists are concerned that biodiversity loss from human impact on natural ecosystems could compromise ecosystem stability. A relationship between diversity and stability was proposed by MacArthur [MacArthur, R.H., 1955. Fluctuation of animal populations and a measure of community stability. Ecology 36, 533–536.]. Current thinking (for example, McCann, K., 2000. The diversity–stability debate. Nature 405, 228–233.) acknowledges that interaction pattern among species, rather than species richness per se, is one element of this relationship. Dunne et al. [Dunne, J.A., Williams, R.J., Martinez, N.D., 2002a. Network structure and biodiversity loss in food webs: robustness increases with connectance. Ecol. Lett. 5, 558–567.] showed that the robustness of 16 food webs is correlated with their connectance. Connectance is one measure of interaction pattern. Robustness relates to the maintenance of network integrity and so has consequences for stability; the loss of integrity must have ecosystem-wide implications. This paper tests the hypothesis that changes in a food web's connectance indicate changes in its robustness. It concludes that any change in connectance with species loss, but especially large, negative changes, constitutes a decrease in robustness. Estimation of the change in connectance could support interpretation of monitoring data on species composition, acting as an indicator of food web robustness and, indirectly, of ecosystem stability. It could assist managers to understand the implications of biodiversity loss caused by human intervention in ecosystems, and could assist either choice of intervention or amelioration of impacts.