Structural Degradation in Mediterranean Sea Food Webs: Testing Ecological Hypotheses Using Stochastic and Mass-Balance Modelling

Human-mediated disturbances such as fishing, habitat modification, and pollution have resulted in significant shifts in species composition and abundance in marine ecosystems which translate into degradation of food-web structure. Here, we used a comparative ecological modelling approach and data from two food webs (North-Central Adriatic and South Catalan Sea) and two time periods (mid-late 1970s and 1990s) in the Mediterranean Sea to evaluate how changes in species composition and biomass have affected food-web properties and the extent of ecosystem degradation. We assembled species lists and ecological information for both regions and time periods into stochastic structural and mass-balance food-web models, and compared the outcomes of 22 food-web properties. Our results show strong similarities in structural food-web properties between the North-Central Adriatic and South Catalan Seas indicating similar ecosystem structure and levels of ecological degradation between regions and time periods. In contrast, a comparison with other published marine food webs (Caribbean, Benguela, and US continental shelf) suggested that Mediterranean webs are in an advanced state of ecological degradation. This was reflected by lower trophic height, linkage density, connectance, omnivory, species involved in looping, trophic chain length and fraction of biomass at higher trophic levels, as well as higher generality and fraction of biomass at lower trophic levels. An analysis of robustness to simulated species extinction revealed lower robustness to species removals in Mediterranean webs and corroborated their advanced state of degradation. Importantly, the two modelling approaches used delivered comparable results suggesting that they both capture fundamental information about how food webs are structured. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Food Webs in the Human Body: Linking Ecological Theory to Viral Dynamics

The dynamics of in-host infections are central to predicting the progression of natural infections and the effectiveness of drugs or vaccines, however, they are not well understood. Here, we apply food web theory to in-host disease networks of the human body that are structured similarly to food web models that treat both predation and competition simultaneously. We show that in-host trade-offs, an under-studied aspect of disease ecology, are fundamental to understanding the outcomes of competing viral strains under differential immune responses. Further, and importantly, our analysis shows that the outcome of competition between virulent and non-virulent strains can be highly contingent on the abiotic conditions prevailing in the human body. These results suggest the alarming idea that even subtle behavioral changes that alter the human body (e.g. weight gain, smoking) may switch the environmental conditions in a manner that suddenly allows a virulent strain to dominate and replace less virulent strains. These ecological results therefore cast new light on the control of disease in the human body, and highlight the importance of longitudinal empirical studies across host variation gradients, as well as, of studies focused on delineating life history trade-offs within hosts.     

Food Webs in Relation to Variation in the Environment and Species Assemblage: A Multivariate Approach

The abiotic environment has strong influences on the growth, survival, behavior, and ecology of aquatic organisms. Biotic interactions and species life histories interact with abiotic factors to structure the food web. One measure of food-web structure is food-chain length. Several hypotheses predict a linear relationship between one environmental variable (e.g., disturbance or ecosystem size) and food-chain length. However, many abiotic and biotic variables interact in diverse ways to structure a community, and may affect other measures of food web structure besides food-chain length. This study took a multivariate approach to test the influence of several important environmental variables on four food-web characteristics measured in nine ponds along a hydroperiod gradient over two years. This approach allowed for testing the ecosystem size and dynamic constraints hypotheses while in context of other possibly interacting environmental variables. The relationship between amphibian and invertebrate communities and pond habitat variables was assessed to understand the underlying food-web structure. Hydroperiod and pond area had a strong influence on amphibian and invertebrate communities, trophic diversity and δ¹⁵N range. The range in δ¹³C values responded strongly to dissolved oxygen. Food-chain length responded to multiple environmental variables. Invertebrate and amphibian communities were structured by pond hydroperiod which in turn influenced the trophic diversity of the food web. The results of this study suggest food-chain length is influenced by environmental variation and species assemblage and that a multivariate approach may allow us to better understand the dynamics within and across aquatic food webs.     

Landscape-Scale Hydrologic Characteristics Differentiate Patterns of Carbon Flow in Large-River Food Webs

Efforts to conserve, restore, or otherwise manage large rivers and the services they provide are hindered by limited understanding of the functional dynamics of these systems. This shortcoming is especially evident with regard to trophic structure and energy flow. We used natural abundances of carbon and nitrogen isotopes to examine patterns of material flow in ten large-river food webs characterized by different landscape-scale hydrologic characteristics (low-gradient river, high-gradient river, river stretches downstream of reservoirs, and reservoirs), and tested predictions from three ecosystem concepts commonly applied to large-rivers: The River Continuum Concept, The Flood Pulse Concept and the Riverine Productivity Model. Carbon derived from aquatic C₃ plants and phytoplankton were the dominant energy sources supporting secondary consumers across the ten large-river food webs examined, but relative contributions differed significantly among landscape types. For low-gradient river food webs, aquatic C₃ plants were the principal carbon source, contributing as much as 80% of carbon assimilated by top consumers, with phytoplankton secondarily important. The estimated relative importance of phytoplankton was greatest for food webs of reservoirs and river stretches downriver from impoundments, although aquatic C₃ plants contributed similar amounts in both landscape types. Highest 99th percentile source contribution estimates for C₄ plants and filamentous algae (both approximately 40%) were observed for high-gradient river food webs. Our results for low-gradient rivers supported predictions of the Flood Pulse Concept, whereas results for the three other landscape types supported the Riverine Productivity Model to varying degrees. Incorporation of landscape-scale hydrologic or geomorphic characteristics, such as river slope or floodplain width, may promote integration of fluvial ecosystem concepts. Expanding these models to include hydrologically impacted landscapes should lead to a more holistic understanding of ecosystem processes in large-river systems.     

Folk Food Webs and the Role of Praxis in Substantive Ecological Knowledge

In the second half of the 20th century, investigations of indigenous environmental knowledge have been the subject of broader anthropological debates over how knowledge and experience are formed. Many such approaches have focused on environmental nomenclature and taxonomy, or what Roy Ellen has called “formal lexical knowledge” (1999). Such knowledge is readily available to an ethnographer and also more easily transmitted through language between subjects. These characteristics of formal lexical knowledge have led to considerable attention given to differences in environmental knowledge between cultures and have possibly resulted in the inflation of the efficacy of language in forming knowledge. However, if a different form of environmental knowledge is examined are there differences that emerge within communities and other processes beyond symbolic systems that shape knowledge? To address these questions, individuals in two Balinese agricultural communities were asked to construct food webs by linking photos of plant and animal species according to ecological interaction. The results showed significant variation in subjects’ knowledge by gender, which corresponds to labor experience in Balinese wet rice agricultural systems. By shifting attention toward emic models of ecological interactions, this article attempts to demonstrate (1) that environmental knowledge differs within a single community; and, (2) the role of labor experience or praxis has in forming environmental knowledge.     

Longer and Less Overlapping Food Webs in Anthropogenically Disturbed Marine Ecosystems: Confirmations from the Past

The human exploitation of marine resources is characterised by the preferential removal of the largest species. Although this is expected to modify the structure of food webs, we have a relatively poor understanding of the potential consequences of such alteration. Here, we take advantage of a collection of ancient consumer tissues, using stable isotope analysis and SIBER to assess changes in the structure of coastal marine food webs in the South-western Atlantic through the second half of the Holocene as a result of the sequential exploitation of marine resources by hunter-gatherers, western sealers and modern fishermen. Samples were collected from shell middens and museums. Shells of both modern and archaeological intertidal herbivorous molluscs were used to reconstruct changes in the stable isotopic baseline, while modern and archaeological bones of the South American sea lion Otaria flavescens, South American fur seal Arctocephalus australis and Magellanic penguin Spheniscus magellanicus were used to analyse changes in the structure of the community of top predators. We found that ancient food webs were shorter, more redundant and more overlapping than current ones, both in northern-central Patagonia and southern Patagonia. These surprising results may be best explained by the huge impact of western sealing on pinnipeds during the fur trade period, rather than the impact of fishing on fish populations. As a consequence, the populations of pinnipeds at the end of the sealing period were likely well below the ecosystem''s carrying capacity, which resulted in a release of intraspecific competition and a shift towards larger and higher trophic level prey. This in turn led to longer and less overlapping food webs.     

Protozoa in Planktonic Food Webs1,2

Protozoa are now being recognized as important members of planktonic food webs. This is due to the inclusion of microbial links in our paradigm of trophic relationships. Heterotrophic microflagellates and ciliates are major grazers of bacteria. They can stimulate production through nutrient recycling and can transform microbial production into larger particles, which are then available for macroconsumers. In this paper we add new groups, the small (< 20 μm) ciliates and myxotrophic flagellates, to the planktonic food web.     

Modelling Size Structured Food Webs Using a Modified Niche Model with Two Predator Traits

The structure of food webs is frequently described using phenomenological stochastic models. A prominent example, the niche model, was found to produce artificial food webs resembling real food webs according to a range of summary statistics. However, the size structure of food webs generated by the niche model and real food webs has not yet been rigorously compared. To fill this void, I use a body mass based version of the niche model and compare prey-predator body mass allometry and predator-prey body mass ratios predicted by the model to empirical data. The results show that the model predicts weaker size structure than observed in many real food webs. I introduce a modified version of the niche model which allows to control the strength of size-dependence of predator-prey links. In this model, optimal prey body mass depends allometrically on predator body mass and on a second trait, such as foraging mode. These empirically motivated extensions of the model allow to represent size structure of real food webs realistically and can be used to generate artificial food webs varying in several aspects of size structure in a controlled way. Hence, by explicitly including the role of species traits, this model provides new opportunities for simulating the consequences of size structure for food web dynamics and stability.     

Industrial Ecosystems and Food Webs: An Expansion and Update of Existing Data for Eco‐Industrial Parks and Understanding the Ecological Food Webs They Wish to Mimic

Cyclical industrial networks are becoming highly desirable for their efficient use of resources and capital. Progress toward this ideal can be enhanced by mimicking the structure of naturally sustainable ecological food webs (FWs). The structures of cyclic industrial networks, sometimes known as eco‐industrial parks (EIPs), are compared to FWs using a variety of important structural ecological parameters. This comparison uses a comprehensive data set of 144 FWs that provides a more ecologically correct understanding of how FWs are organized than previous efforts. In conjunction, an expanded data set of 48 EIPs gives new insights into similarities and differences between the two network types. The new information shows that, at best, current EIPs are most similar to those FWs that lack the components that create a biologically desirable cyclical structure. We propose that FWs collected from 1993 onward should be used in comparisons with EIPs, given that these networks are much more likely to include important network functions that directly affect the structure. We also propose that the metrics used in an ecological analysis of EIPs be calculated from an FW matrix, as opposed to a community matrix, which, to this point, has been widely used. These new insights into the design of ecologically inspired industrial networks clarify the path toward superior material and energy cycling for environmental and financial success.     

Marine Cyanophages Demonstrate Biogeographic Patterns throughout the Global Ocean

Myoviruses and podoviruses that infect cyanobacteria are the two major groups of marine cyanophages, but little is known of how their phylogenetic lineages are distributed in different habitats. In this study, we analyzed the phylogenetic relationships of cyanopodoviruses and cyanomyoviruses based on the existing genomes. The 28 cyanomyoviruses were classified into four clusters (I to IV), and 19 of the 20 cyanopodoviruses were classified into two clusters, MPP-A and MPP-B, with four subclusters within cluster MPP-B. These genomes were used to recruit cyanophage-like fragments from microbial and viral metagenomes to estimate the relative abundances of these cyanophage lineages. Our results showed that cyanopodoviruses and cyanomyoviruses are both abundant in various marine environments and that clusters MPP-B, II and III appear to be the most dominant lineages. Cyanopodoviruses and cluster I and IV cyanomyoviruses exhibited habitat-related variability in their relative levels of abundance, while cluster II and III cyanomyoviruses appeared to be consistently dominant in various habitats. Multivariate analyses showed that reads that mapped to Synechococcus phages and Prochlorococcus phages had distinct distribution patterns that were significantly correlated to those of Synechococcus and Prochlorococcus, respectively. The Mantel test also revealed a strong correlation between the community compositions of cyanophages and picocyanobacteria. Given that cyanomyoviruses tend to have a broad host range and some can cross-infect Synechococcus and Prochlorococcus, while cyanopodoviruses are commonly host specific, the observation that their community compositions both correlated significantly with that of picocyanobacteria was unexpected. Although cyanomyoviruses and cyanopodoviruses differ in host specificity, their biogeographic distributions are likely both constrained by the picocyanobacterial community.     

The Role of Pre-Existing Disturbances in the Effect of Marine Reserves on Coastal Ecosystems: A Modelling Approach

We have used an end-to-end ecosystem model to explore responses over 30 years to coastal no-take reserves covering up to 6% of the fifty thousand square kilometres of continental shelf and slope off the coast of New South Wales (Australia). The model is based on the Atlantis framework, which includes a deterministic, spatially resolved three-dimensional biophysical model that tracks nutrient flows through key biological groups, as well as extraction by a range of fisheries. The model results support previous empirical studies in finding clear benefits of reserves to top predators such as sharks and rays throughout the region, while also showing how many of their major prey groups (including commercial species) experienced significant declines. It was found that the net impact of marine reserves was dependent on the pre-existing levels of disturbance (i.e. fishing pressure), and to a lesser extent on the size of the marine reserves. The high fishing scenario resulted in a strongly perturbed system, where the introduction of marine reserves had clear and mostly direct effects on biomass and functional biodiversity. However, under the lower fishing pressure scenario, the introduction of marine reserves caused both direct positive effects, mainly on shark groups, and indirect negative effects through trophic cascades. Our study illustrates the need to carefully align the design and implementation of marine reserves with policy and management objectives. Trade-offs may exist not only between fisheries and conservation objectives, but also among conservation objectives.     

Tracing Consumer-Derived Nitrogen in Riverine Food Webs

The flux of consumer-derived nutrients is recognized as an important ecosystem process, yet few studies have quantified the impact of these fluxes on freshwater ecosystems. The high abundance of bivalves in both marine and freshwater suggests that bivalves can exert large effects on aquatic food webs. The objective of our study was to determine the importance of unionid mussel-derived nitrogen (MDN) to the food web. We used a stable isotope tracer approach in conjunction with nutrient uptake and excretion experiments. We fed mussels (Lampsilis siliquiodea, n = 249) a ¹⁵N-enriched algal diet and placed them into a N-limited stream for 63 days. Mussel hemolymph was non-lethally sampled over the course of the experiment to measure tissue turnover of δ¹⁵N and excretion experiments were done to model the amount of N mussels provided in comparison to stream N uptake demand. Multiple food web pools were sampled twice prior and five times following the mussel addition to trace the ¹⁵N through the food web. Our mussel excretion rates in comparison to areal uptake demand suggested that mussel excretion can account for 40% of the total N demand in this stream. Our enrichment showed that MDN was entering the food web and supplied up to 19% of the N in specific compartments of the food web near the mussel bed. When scaled to a natural mussel aggregation, our results suggest up to 74% of N in the food web may be mussel-derived. Our results show that N supplied by mussels can be an important nutrient subsidy that provides food web support.