Integrating ecosystem engineering and food webs

Ecosystem engineering, the physical modification of the environment by organisms, is a common and often influential process whose significance to food web structure and dynamics is largely unknown. In the light of recent calls to expand food web studies to include non‐trophic interactions, we explore how we might best integrate ecosystem engineering and food webs. We provide rationales justifying their integration and present a provisional framework identifying how ecosystem engineering can affect the nodes and links of food webs and overall organization; how trophic interactions with the engineer can affect the engineering; and how feedbacks between engineering and trophic interactions can affect food web structure and dynamics. We use a simple integrative food chain model to illustrate how feedbacks between the engineer and the food web can alter 1) engineering effects on food web dynamics, and 2) food web responses to extrinsic environmental perturbations. We identify four general challenges to integration that we argue can readily be met, and call for studies that can achieve this integration and help pave the way to a more general understanding of interaction webs in nature. Synthesis All species are affected by their physical environment. Because ecosystem engineering species modify the physical environment and belong to food webs, such species are potentially one of the most important bridges between the trophic and non‐trophic. We examine how to integrate the so far, largely independent research areas of ecosystem engineering and food webs. We present a conceptual framework for understanding how engineering can affect food webs and vice versa, and how feedbacks between the two alter ecosystem dynamics. With appropriate empirical studies and models, integration is achievable, paving the way to a more general understanding of interaction webs in nature.     

Exploring the temporal variability of a food web using long‐term biomonitoring data

Ecological communities are constantly being reshaped in the face of environmental change and anthropogenic pressures. Yet, how food webs change over time remains poorly understood. Food web science is characterized by a trade‐off between complexity (in terms of the number of species and feeding links) and dynamics. Topological analysis can use complex, highly resolved empirical food web models to explore the architecture of feeding interactions but is limited to a static view, whereas ecosystem models can be dynamic but use highly aggregated food webs. Here, we explore the temporal dynamics of a highly resolved empirical food web over a time period of 18 years, using the German Bight fish and benthic epifauna community as our case study. We relied on long‐term monitoring ecosystem surveys (from 1998 to 2015) to build a metaweb, i.e. the meta food web containing all species recorded over the time span of our study. We then combined time series of species abundances with topological network analysis to construct annual food web snapshots. We developed a new approach, ‘node‐weighted’ food web metrics by including species abundances to represent the temporal dynamics of food web structure, focusing on generality and vulnerability. Our results suggest that structural food web properties change through time; however, binary food web structural properties may not be as temporally variable as the underlying changes in species composition. Further, the node‐weighted metrics enabled us to detect that food web structure was influenced by changes in species composition during the first half of the time series and more strongly by changes in species dominance during the second half. Our results demonstrate how ecosystem surveys can be used to monitor temporal changes in food web structure, which are important ecosystem indicators for building marine management and conservation plans.     

A cascade of evolutionary change alters consumer-resource dynamics and ecosystem function

It is becoming increasingly clear that intraspecific evolutionary divergence influences the properties of populations, communities and ecosystems. The different ecological impacts of phenotypes and genotypes may alter selection on many species and promote a cascade of ecological and evolutionary change throughout the food web. Theory predicts that evolutionary interactions across trophic levels may contribute to hypothesized feedbacks between ecology and evolution. However, the importance of 'cascading evolutionary change' in a natural setting is unknown. In lakes in Connecticut, USA, variation in migratory behaviour and feeding morphology of a fish predator, the alewife (Alosa pseudoharengus), drives life-history evolution in a species of zooplankton prey (Daphnia ambigua). Here we evaluated the reciprocal impacts of Daphnia evolution on ecological processes in laboratory mesocosms. We show that life-history evolution in Daphnia facilitates divergence in rates of population growth, which in turn significantly alters consumer-resource dynamics and ecosystem function. These experimental results parallel trends observed in lakes. Such results argue that a cascade of evolutionary change, which has occurred over contemporary timescales, alters community and ecosystem processes.     

The significance of partial migration for food web and ecosystem dynamics

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.     

The role of fish life histories in allometrically scaled food‐web dynamics

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A decade of seasonal dynamics and co-occurrences within freshwater bacterioplankton communities from eutrophic Lake Mendota,WI, USA

With an unprecedented decade-long time series from a temperate eutrophic lake, we analyzed bacterial and environmental co-occurrence networks to gain insight into seasonal dynamics at the community level. We found that (1) bacterial co-occurrence networks were non-random, (2) season explained the network complexity and (3) co-occurrence network complexity was negatively correlated with the underlying community diversity across different seasons. Network complexity was not related to the variance of associated environmental factors. Temperature and productivity may drive changes in diversity across seasons in temperate aquatic systems, much as they control diversity across latitude. While the implications of bacterioplankton network structure on ecosystem function are still largely unknown, network analysis, in conjunction with traditional multivariate techniques, continues to increase our understanding of bacterioplankton temporal dynamics.     

Unravelling the complex structure of forest soil food webs: higher omnivory and more trophic levels

Food web topologies depict the community structure as distributions of feeding interactions across populations. Although the soil ecosystem provides important functions for aboveground ecosystems, data on complex soil food webs is notoriously scarce, most likely due to the difficulty of sampling and characterizing the system. To fill this gap we assembled the complex food webs of 48 forest soil communities. The food webs comprise 89 to 168 taxa and 729 to 3344 feeding interactions. The feeding links were established by combining several molecular methods (stable isotope, fatty acid and molecular gut content analyses) with feeding trials and literature data. First, we addressed whether soil food webs (n = 48) differ significantly from those of other ecosystem types (aquatic and terrestrial aboveground, n = 77) by comparing 22 food web parameters. We found that our soil food webs are characterized by many omnivorous and cannibalistic species, more trophic chains and intraguild‐predation motifs than other food webs and high average and maximum trophic levels. Despite this, we also found that soil food webs have a similar connectance as other ecosystems, but interestingly a higher link density and clustering coefficient. These differences in network structure to other ecosystem types may be a result of ecosystem specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food‐web level. In a second analysis of land‐use effects, we found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. Overall, our study has unravelled some systematic structures of soil food‐webs, which extends our mechanistic understanding how environmental characteristics of the soil ecosystem determine patterns at the community level.     

食物网稳定性机制研究进展:一个基于等级系统的框架

食物网理论沟通了群落生态学和生态系统生态学,将生物多样性和生态系统功能的研究统一起来,是理解生态系统运作机制的关键。自从1973年Robert May的经典研究引发著名的"复杂性-稳定性"论辩之后,人们认识到食物网的稳定性是其结构维持、功能发挥和动态演化的一个重要前提,并开始了对食物网稳定性机制的探索。早期研究主要关注只包含拓扑关系的定性食物网,但后来人们逐渐认识到相互作用强度的重要性,并提出了诸如自限性、弱相互作用、适应性捕食等一系列机制。本文系统梳理了过往研究中模块层面的各类稳定性机制和全网层面对各模块的整合机制,从而清晰地展示了"模块-全网"双层框架的全貌。通过在其基础上的扩展,进而提出了一个基于等级系统的食物网稳定性框架,并从动力学和能量学角度,对各层级内部的稳定性机制以及层级之间的关系进行了探讨,以期为建立普适的食物网稳定性理论提供一些思路。未来的研究方向包括：①将稳定性机制的研究从食物网扩展到更一般的生态网络;②综合考虑生物物理要素、动力学稳定性、系统对能流功率的追求、环境的平稳程度、演化历史等影响因素,从而得到关于食物网结构和动态的更为深刻的认识。     

Language as an evolving word web.

Human language may be described as a complex network of linked words. In such a treatment, each distinct word in language is a vertex of this web, and interacting words in sentences are connected by edges. The empirical distribution of the number of connections of words in this network is of a peculiar form that includes two pronounced power-law regions. Here we propose a theory of the evolution of language, which treats language as a self-organizing network of interacting words. In the framework of this concept, we completely describe the observed word web structure without any fitting. We show that the two regimes in the distribution naturally emerge from the evolutionary dynamics of the word web. It follows from our theory that the size of the core part of language, the 'kernel lexicon', does not vary as language evolves.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Climate change alters the structure of arctic marine food webs due to poleward shifts of boreal generalists

Climate-driven poleward shifts, leading to changes in species composition and relative abundances, have been recently documented in the Arctic. Among the fastest moving species are boreal generalist fish which are expected to affect arctic marine food web structure and ecosystem functioning substantially. Here, we address structural changes at the food web level induced by poleward shifts via topological network analysis of highly resolved boreal and arctic food webs of the Barents Sea. We detected considerable differences in structural properties and link configuration between the boreal and the arctic food webs, the latter being more modular and less connected. We found that a main characteristic of the boreal fish moving poleward into the arctic region of the Barents Sea is high generalism, a property that increases connectance and reduces modularity in the arctic marine food web. Our results reveal that habitats form natural boundaries for food web modules, and that generalists play an important functional role in coupling pelagic and benthic modules. We posit that these habitat couplers have the potential to promote the transfer of energy and matter between habitats, but also the spread of pertubations, thereby changing arctic marine food web structure considerably with implications for ecosystem dynamics and functioning.     

Novel feeding interactions amplify the impact of species redistribution on an Arctic food web

Species are redistributing globally in response to climate warming, impacting ecosystem functions and services. In the Barents Sea, poleward expansion of boreal species and a decreased abundance of Arctic species are causing a rapid borealization of the Arctic communities. This borealization might have profound consequences on the Arctic food web by creating novel feeding interactions between previously non co‐occurring species. An early identification of new feeding links is crucial to predict their ecological impact. However, detection by traditional approaches, including stomach content and isotope analyses, although fundamental, cannot cope with the speed of change observed in the region, nor with the urgency of understanding the consequences of species redistribution for the marine ecosystem. In this study, we used an extensive food web (metaweb) with nearly 2,500 documented feeding links between 239 taxa coupled with a trait data set to predict novel feeding interactions and to quantify their potential impact on Arctic food web structure. We found that feeding interactions are largely determined by the body size of interacting species, although species foraging habitat and metabolic type are also important predictors. Further, we found that all boreal species will have at least one potential resource in the Arctic region should they redistribute therein. During 2014–2017, 11 boreal species were observed in the Arctic region of the Barents Sea. These incoming species, which are all generalists, change the structural properties of the Arctic food web by increasing connectance and decreasing modularity. In addition, these boreal species are predicted to initiate novel feeding interactions with the Arctic residents, which might amplify their impact on Arctic food web structure affecting ecosystem functioning and vulnerability. Under the ongoing species redistribution caused by environmental change, we propose merging a trait‐based approach with ecological network analysis to efficiently predict the impacts of range‐shifting species on food webs.     

Accommodating environmental variation in population models: metaphysiological biomass loss accounting

1. There is a pressing need for population models that can reliably predict responses to changing environmental conditions and diagnose the causes of variation in abundance in space as well as through time. In this 'how to' article, it is outlined how standard population models can be modified to accommodate environmental variation in a heuristically conducive way. This approach is based on metaphysiological modelling concepts linking populations within food web contexts and underlying behaviour governing resource selection. Using population biomass as the currency, population changes can be considered at fine temporal scales taking into account seasonal variation. Density feedbacks are generated through the seasonal depression of resources even in the absence of interference competition. 2. Examples described include (i) metaphysiological modifications of Lotka-Volterra equations for coupled consumer-resource dynamics, accommodating seasonal variation in resource quality as well as availability, resource-dependent mortality and additive predation, (ii) spatial variation in habitat suitability evident from the population abundance attained, taking into account resource heterogeneity and consumer choice using empirical data, (iii) accommodating population structure through the variable sensitivity of life-history stages to resource deficiencies, affecting susceptibility to oscillatory dynamics and (iv) expansion of density-dependent equations to accommodate various biomass losses reducing population growth rate below its potential, including reductions in reproductive outputs. Supporting computational code and parameter values are provided. 3. The essential features of metaphysiological population models include (i) the biomass currency enabling within-year dynamics to be represented appropriately, (ii) distinguishing various processes reducing population growth below its potential, (iii) structural consistency in the representation of interacting populations and (iv) capacity to accommodate environmental variation in space as well as through time. Biomass dynamics provide a common currency linking behavioural, population and food web ecology. 4. Metaphysiological biomass loss accounting provides a conceptual framework more conducive for projecting and interpreting the population consequences of climatic shifts and human transformations of habitats than standard modelling approaches.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

Interplay between the paradox of enrichment and nutrient cycling in food webs

Nutrient cycling is fundamental to ecosystem functioning. Despite recent major advances in the understanding of complex food web dynamics, food web models have so far generally ignored nutrient cycling. However, nutrient cycling is expected to strongly impact food web stability and functioning. To make up for this gap, we built an allometric and size structured food web model including nutrient cycling. By releasing mineral nutrients, recycling increases the availability of limiting resources for primary producers and links each trophic level to the bottom of food webs. We found that nutrient cycling can provide a significant part of the total nutrient supply of the food web, leading to a strong enrichment effect that promotes species persistence in nutrient poor ecosystems but leads to a paradox of enrichment at high nutrient inputs. The presence of recycling loops linking each trophic level to the basal resources weakly affects species biomass temporal variability in the food web. Recycling loops tend to slightly dampen the destabilising effect of nutrient enrichment on consumer temporal variability while they have opposite effects for primary producers. By considering nutrient cycling, this new model improves our understanding of the response of food webs to nutrient availability and opens perspectives to better link studies on food web dynamics and ecosystem functioning.     

The consequences of mass mortality events for the structure and dynamics of biological communities

Mass mortality events (MMEs) are rapidly occurring, substantial population losses that transpire within a short time interval relative to the generation time of the affected organism. Previous work has established that MMEs appear to be increasing in frequency and magnitude; however, currently, there is little understanding of the consequences of MMEs for biological communities. Here, we use theory and empirical data from observed MMEs to understand how MMEs impact the structure and dynamics of communities. To do so, we build upon existing resource pulse and trophic cascade theory to show that MMEs both share similarities and diverge from these ecological phenomena, producing distinct short‐ and long‐term impacts by jointly altering the effects of species interactions across trophic levels and providing an influx of resources from decaying biomass. Second, we investigate how the magnitude of MMEs, trophic level of the impacted species, overall food web structure and ecosystem type may mediate the resulting ecological response. Third, we compare the understanding gained by our models to existing observational data on MMEs. Our synthesis, offers an empirical path forward for understanding MMEs through experimentation and improved observational data collection. While complex, resolving the consequences of MMEs should be a high research priority due to their role in determining how ecological systems respond to environmental change driven by rare events.     

Communities at the extreme: Aquatic food webs in desert landscapes

Studying food webs across contrasting abiotic conditions is an important tool in understanding how environmental variability impacts community structure and ecosystem dynamics. The study of extreme environments provides insight into community‐wide level responses to environmental pressures with relevance to the future management of aquatic ecosystems. In the western Lake Eyre Basin of arid Australia, there are two characteristic and contrasting aquatic habitats: springs and rivers. Permanent isolated Great Artesian Basin springs represent hydrologically persistent environments in an arid desert landscape. In contrast, hydrologically variable river waterholes are ephemeral in space and time. We comprehensively sampled aquatic assemblages in contrasting ecosystem types to assess patterns in community composition and to quantify food web attributes with stable isotopes. Springs and rivers were found to have markedly different invertebrate communities, with rivers dominated by more dispersive species and springs associated with species that show high local endemism. Qualitative assessment of basal resources shows autochthonous carbon appears to be a key basal resource in both types of habitat, although the particular sources differed between habitats. Food‐web variables such as trophic length, trophic breadth, and community isotopic niche size were relatively similar in the two habitat types. The basis for the similarity in food‐web structure despite differences in community composition appears to be broader isotopic niches for predatory invertebrates and fish in springs as compared with rivers. In contrast to published theory, our findings suggest that the food webs of the hydrologically variable river sites may show less dietary generalization and more compact food‐web modules than in springs.