Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) are used to study the trophic structure of food web in the Yellow Sea and East China Sea ecosystem. The trophic continuum of pelagic food web from phytoplankton to top preyer was elementarily established, and a trophic structure diagram in the Yellow Sea and East China Sea was outlined in combination with carbon isotopic data of benthic organisms, which is basically consistent with and makes some improvements on the simplified Yellow Sea food web and the trophic structure diagram drawn based on the biomass of main resource population during 1985–1986. This result indicates that the stable isotope method is a potential useful means for further studying the complete marine food web trophic continuum from viruses to top predators and food web stability.     

Trophic relationships: integrating meiofauna into a realistic benthic food web

• 1 This paper summarises the most important contributions on trophic relationships of lotic meiofauna. In contrast to marine research, the few quantitative studies of the freshwater meiobenthos have shown that these invertebrates not only take up particulate/fine organic matter, but also dissolved organic substances attached to organic particles. In lotic ecosystems, further estimates of grazing rate and bacterial/algal ingestion rate are needed, particularly in situ measurements.
• 2 The effects of macroinvertebrate predators upon meiofauna are still under debate. Depending on the type of experiments (laboratory vs. field) it seems that macrofauna may or may not affect meiofauna. Field samples and analyses of gut contents of larval tanypod chironomids have shown that the impact upon meiofauna was low and larvae were nonselective predators. Predation amounted to 2.2% of the combined prey density and prey consumption averaged 1.3 individuals per predator individual per year.
• 3 Adding taxonomic resolution by including the meiofaunal component within lotic food webs distinctly increases the number of total species and, as a consequence, changes food web statistics. Webs that included meiofauna revealed that these metazoans contributed substantially to the percentage of intermediate species (species with predators and prey). The resolution of dietary analyses of major consumers of macro‐ and meiobenthos showed that many stream invertebrates feed on meiofauna.

     

Food web structure variability in the surface layer, at a fixed station influenced by the North Western Mediterranean Current

The structure of the planktonic community and the influence of mesozooplankton migration on the microbial food web were investigated during six diel studies from June 92 to June 93 in the surface waters of a station in the North-Western Mediterranean Sea. Each diel study consisted of sampling at 5 and 40 m every 3 h over 24 h. Most of the times diel cycles did not show any convincing diel patterns in any of the variables studied. Clear zooplankton migration was evident in only two diel studies.The ratio of heterotrophic/autotrophic biomasses varied from 0.68 to 3.0, with a strong dominance of the heterotrophic biomass under oligotrophic conditions. Differences in food web structure were probably related to the influences of coastal water and the North-Western Mediterranean Current. Thus we found that the planktonic food web variability relatable to hydrodynamic variability, to be greater than diel variability. However, very large differences in food web structure among dates were evident. For example proportion of Chl a found in the <10 µm fraction varied from 18 to 96%.     

Beach morphology and food web structure: comparison of an eroding and an accreting sandy shore in the North Sea

Food web components and inorganic nutrients were studied on two sandy shores of the adjacent barrier islands of Sylt and R?m? in the North Sea, differing in morphodynamics. Implications of high and low wave energy on the food web structure were assessed. The Sylt shore represents a dynamic intermediate beach type, while the R?m? shore is morphologically stable and dissipative. On the steep-profiled, coarse-grained Sylt shore, strong hydrodynamics resulted in erosion and high fluxes of organic material through the beach, but prevented any storage of food sources. In contrast, the flat-profiled, fine-grained R?m? shore, with low wave energy and accretion, accumulated organic carbon from surf waters. At Sylt, oxic nutrient regeneration prevailed, while anoxic mineralization was more important at R?m?. Macrofauna on the Sylt shore was impoverished compared with the community at R?m?. Correspondingly, abundances of epibenthic predators such as shrimps, crabs, fish, and shorebirds were also lower at Sylt. Meiofauna was abundant on both shores, but differed in taxonomic composition. Several major taxa were represented in fairly equal proportions of individual numbers on the well-oxygenated Sylt shore, while nematodes strongly dominated the assemblage at R?m?. Thus, on cold-temperate, highly dynamic intermediate shores with high wave energy and subject to erosion, the "small food web" dominates. Organisms are agile and quickly exploit fresh organic material. Larger organisms and nematodes abound under stable, dissipative and accreting shore conditions, where some food materials may accumulate and zoomass builds up to support numerous visitors from higher trophic levels. Electronic Publication     

Fit, efficiency, and biology: some thoughts on judging food web models

Revealing the processes that determine who eats whom, and thereby the structure of food webs, is a long running challenge in ecological research. Recent advances include development of new methods for measuring fit of models to observed food web data, and thereby testing which are the ‘best’ food web models. The best model could be considered the most efficient with relatively few parameters and high explanatory power. Another recent advance involves adding some additional biology to food web models in the form of foraging theory based on maximisation of energy intake as the predictor of species' diets in food webs. While it is interesting to compare efficiency among food web models, we believe that such comparisons at least should be interpreted with caution, since they do not account for any differences in motivation, formulation, and potential that might also exist among models. Furthermore, we see an important but somewhat neglected role for experimental tests of models of food web structure.     

Trophic interactions within the microbial food web in the South China Sea revealed by size-fractionation method

To define nanoflagellate-bacteria interactions and potential trophic levels within the microbial food web in the oligotrophic South China Sea, we conducted fourteen size-fractionation experiments in which seawater was filtered through 1, 2, 5, 10, 20, 60, and 200 μm membranes or meshes and the growth of four groups of picoplankton, Prochlorococcus, Synechococcus, high DNA heterotrophic bacteria, and low DNA heterotrophic bacteria were monitored in each filtrate after 24 hours of incubation. Removing grazers by filtration would relieve the grazing pressure on lower trophic levels which finally influenced the net growth rates of picoplankton. The growth patterns of Prochlorococcus and Synechococcus were similar, with higher growth rates in the < 1 μm or < 2 μm treatments, a second peak in the < 10 μm treatments and often a third peak in the < 200 μm treatments. The net growth rates of low DNA heterotrophic bacteria were little influenced by size-fractionation. Due to a subgroup of high DNA heterotrophic bacteria with larger size and higher DNA content which appeared to resist the grazing by < 5 μm nanoflagellates, the net growth rates of high DNA heterotrophic bacteria were higher in the < 2 μm or < 5 μm treatments with a second peak in the < 60 μm treatments. A general pattern of five potential trophic levels (< 2 μm, 2-5 μm, 5-10 μm, 10-60 μm, 60-200 μm) was revealed combining all the experiments, confirming the existence of multiple trophic levels within the microbial food web in the oligotrophic South China Sea.     

Marine molluscs and fish as biomarkers of pollution stress in littoral regions of the Red Sea, Mediterranean Sea and North Sea

The intensive development of industry and urban structures along the seashores of the world, as well as the immense increase in marine transportation and other activities, has resulted in the deposition of thousands of new chemicals and organic compounds, endangering the existence of organisms and ecosystems. The conventional single biomarker methods used in ecological assessment studies cannot provide an adequate base for environmental health assessment, management and sustainability planning. The present study uses a set of novel biochemical, physiological, cytogenetic and morphological methods to characterize the state of health of selected molluscs and fish along the shores of the German North Sea, as well as the Israeli Mediterranean and Red Sea. The methods include measurement of activity of multixenobiotic resistance-mediated transporter (MXRtr) and the system of active transport of organic anions (SATOA) as indicators of antixenobiotic defence; glutathione S-transferase (GST) activity as an indicator of biotransformation of xenobiotics; DNA unwinding as a marker of genotoxicity; micronucleus test for clastogenicity; levels of phagocytosis for immunotoxicity; cholinesterase (ChE) activity and level of catecholamines as indicators of neurotoxicity; permeability of external epithelia to anionic hydrophilic probe, intralysosomal accumulation of cationic amphiphilic probe and activity of non-specific esterases as indicators of cell/tissue viability. Complete histopathological examination was used for diagnostics of environmental pathology. The obtained data show that the activity of the defensive pumps, MXRtr and SATOA in the studied organisms was significantly higher in the surface epithelia of molluscs from a polluted site than that of the same species from control, unpolluted stations, providing clear evidence of response to stress. Enhanced frequency of DNA lesions (alkaline and acidic DNA unwinding) and micronucleus-containing cells was significantly higher in samples from polluted sites in comparison to those from the clean sites that exhibited genotoxic and clastogenic activity of the pollutants. In all the studied molluscs a negative correlation was found between the MXRtr levels of activity and the frequency of micronucleus-containing hemocytes. The expression of this was in accordance with the level of pollution. The complete histopathological examination demonstrates significantly higher frequencies of pathological alterations in organs of animals from polluted sites. A strong negative correlation was found between the frequency of these alterations and MXRtr activity in the same specimens. In addition to these parameters, a decrease in the viability was noted in molluscs from the polluted sites, but ChE activities remained similar at most sites. The methods applied in our study unmasked numerous early cryptic responses and negative alterations of health in populations of marine biota sampled from the polluted sites. This demonstrates that genotoxic, clastogenic and pathogenic xenobiotics are present and act in the studied sites and this knowledge can provide a reliable base for consideration for sustainable development. Received: 2 March 1999 / Received in revised form: 2 August 1999 / Accepted: 3 August 1999     

Food-web structure and ecosystem services: insights from the Serengeti

The central organizing theme of this paper is to discuss the dynamics of the Serengeti grassland ecosystem from the perspective of recent developments in food-web theory. The seasonal rainfall patterns that characterize the East African climate create an annually oscillating, large-scale, spatial mosaic of feeding opportunities for the larger ungulates in the Serengeti; this in turn creates a significant annual variation in the food available for their predators. At a smaller spatial scale, periodic fires during the dry season create patches of highly nutritious grazing that are eaten in preference to the surrounding older patches of less palatable vegetation. The species interactions between herbivores and plants, and carnivores and herbivores, are hierarchically nested in the Serengeti food web, with the largest bodied consumers on each trophic level having the broadest diets that include species from a large variety of different habitats in the ecosystem. The different major habitats of the Serengeti are also used in a nested fashion; the highly nutritious forage of the short grass plains is available only to the larger migratory species for a few months each year. The longer grass areas, the woodlands and kopjes (large partially wooded rocky islands in the surrounding mosaic of grassland) contain species that are resident throughout the year; these species often have smaller body size and more specialized diets than the migratory species. Only the larger herbivores and carnivores obtain their nutrition from all the different major habitat types in the ecosystem. The net effect of this is to create a nested hierarchy of subchains of energy flow within the larger Serengeti food web; these flows are seasonally forced by rainfall and operate at different rates in different major branches of the web. The nested structure that couples sequential trophic levels together interacts with annual seasonal variation in the fast and slow chains of nutrient flow in a way that is likely to be central to the stability of the whole web. If the Serengeti is to be successfully conserved as a fully functioning ecosystem, then it is essential that the full diversity of natural habitats be maintained within the greater Serengeti ecosystem. The best way to do this is by controlling the external forces that threaten the boundaries of the ecosystem and by balancing the economic services the park provides between local, national and international needs. I conclude by discussing how the ecosystem services provided by the Serengeti are driven by species on different trophic levels. Tourism provides the largest financial revenue to the national economy, but it could be better organized to provide more sustained revenue to the park. Ultimately, ecotourism needs to be developed in ways that take lessons from the structure of the Serengeti food webs, and in ways that provide tangible benefits to people living around the park while also improving the experience of all visitors.     

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20 m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms. Received in revised form: 7 May 2001 Electronic Publication     

Carbon steady-state model of the planktonic food web of Lake Biwa, Japan

1. A steady‐state model of carbon flows was developed to describe the summer planktonic food web in the surface mixed‐layer of the North Basin in Lake Biwa, Japan. This model synthesised results from numerous studies on the plankton of Lake Biwa. 2. An inverse analysis procedure was used to estimate missing flow values in a manner consistent with known information. Network analysis was applied to characterise emergent properties of the resulting food web. 3. The system strongly relied on flows related to detrital particles. Whereas primary production was mainly by phytoplankton >20 μm, microzooplankton were active and mainly ingested detritus and bacteria. 4. The main emergent property of the system was strong recycling, through either direct ingestion of non‐living material by zooplankton, or ingestion of bacteria after degradation of detritus to release dissolved organic carbon.     

Regime shifts in marine communities: a complex systems perspective on food web dynamics

Species composition and habitats are changing at unprecedented rates in the world''s oceans, potentially causing entire food webs to shift to structurally and functionally different regimes. Despite the severity of these regime shifts, elucidating the precise nature of their underlying processes has remained difficult. We address this challenge with a new analytic approach to detect and assess the relative strength of different driving processes in food webs. Our study draws on complexity theory, and integrates the network-centric exponential random graph modelling (ERGM) framework developed within the social sciences with community ecology. In contrast to previous research, this approach makes clear assumptions of direction of causality and accommodates a dynamic perspective on the emergence of food webs. We apply our approach to analysing food webs of the Baltic Sea before and after a previously reported regime shift. Our results show that the dominant food web processes have remained largely the same, although we detect changes in their magnitudes. The results indicate that the reported regime shift may not be a system-wide shift, but instead involve a limited number of species. Our study emphasizes the importance of community-wide analysis on marine regime shifts and introduces a novel approach to examine food webs.     

Contrasting spatial patterns of taxonomic and functional richness offer insights into potential loss of ecosystem services

Functional and trophic perspectives on patterns of species occurrences have the potential to offer new and interesting insights into a range of spatially explicit problems in ecology and conservation. We present the function–area relationship (FAR) and explore linkages between functional and taxonomic species richness for South African birds. We first used beak morphology to classify a subset of 151 South African bird species into 18 functional groups and calculated both the species–area relationship and the FAR at quarter-degree resolution for South Africa. The relationship between functional and taxonomic richness by cell was quadratic rather than linear, with considerable scatter around the curve. We next looked at the spatial relationships between taxonomic diversity and response diversity (i.e. diversity within functional groups) using an a priori categorization of nearly all South African birds into nine functional groups. The spatial distribution of response richness also showed considerable variation in relation to taxonomic richness. Our results demonstrate a novel approach to linking taxonomic, functional and trophic patterns in space and suggest a way in which conservation planning, which has traditionally had a taxonomic focus, could formally incorporate a more functional and food-web-based approach.     

Out of the North Sea: the Zeeland Ridges Neandertal

In 2001, a portion of human frontal bone was discovered in sediments extracted from the bottom of the North Sea, 15 km off the coast of the Netherlands. The extraction zone is located in the so-called Zeeland Ridges area located at 51°40′ northern latitude and 3°20′ eastern longitude. The specimen was dredged up from sediments containing Late Pleistocene faunal remains and Middle Palaeolithic artefacts, including well-finished small handaxes and Levallois flakes. The details of the supraorbital morphology, as well as the quantitative assessment of the shape of the external surface of the squama using traditional and 3D geometric morphometrics, unambiguously assign the Zeeland Ridges frontal bone to Homo neanderthalensis. Carbon and nitrogen isotopic analysis indicate that the Zeeland Ridges hominin, like other Neandertals, was highly carnivorous and does not show evidence for the consumption of aquatic foods. A lesion on the outer table and diploic layer of the bone in the area of the supratoral sulcus can be interpreted as the result of an intradiploic epidermoid cyst, a type of neoplasm diagnosed for the first time in Neandertal remains. So far, the Zeeland Ridges Neandertal is the first Pleistocene fossil hominin found under seawater and the first recorded in the Netherlands.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

Community assembly, stability and signatures of dynamical constraints on food web structure

To understand the dynamics of natural species communities, a major challenge is to quantify the relationship between their assembly, stability, and underlying food web structure. To this end, two complementary aspects of food web structure can be related to community stability: sign structure, which refers to the distributions of trophic links irrespective of interaction strengths, and interaction strength structure, which refers to the distributions of interaction strengths with or without consideration of sign structure. In this paper, using data from a set of relatively well documented community food webs, I show that natural communities generally exhibit a sign structure that renders their stability sensitive to interaction strengths. Using a Lotka-Volterra type population dynamical model, I then show that in such communities, individual consumer species with high values of a measure of their total biomass acquisition rate, which I term “weighted generality”, tend to undermine community stability. Thus consumer species’ trophic modules (a species and all its resource links) should be “selected” through repeated immigrations and extinctions during assembly into configurations that increase the probability of stable coexistence within the constraints of the community's trophic sign structure. The presence of such constraints can be detected by the incidence and strength of certain non-random structural characteristics. These structural signatures of dynamical constraints are readily measurable, and can be used to gauge the importance of interaction-driven dynamical constraints on communities during and after assembly in natural communities.     

The Importance of Ship Hull Fouling as a Vector of Species Introductions into the North Sea

Ships have long been recognized as a major vector for the introduction of non-native and harmful organisms. From 1992 to 1996 a shipping study was undertaken in Germany, focusing on the fauna transported by ships, to assess the importance of species introductions by international shipping traffic. Ballast water, tank sediment or hull fouling of 186 vessels was sampled. A total of 257 species were identified, ranging from Foraminifera to Teleostei, and 57% of the species sampled were considered to be non-native to the North Sea region, originating from elsewhere in the world including the north eastern Atlantic (west of the English Channel). Non-native species were recorded in 38% of all ballast water samples, 57% of all sediment samples and 96% of all hull samples, indicating that hull fouling is an important vector of introduction. Four species (1.6%) of unknown origin (cryptogenic species) were identified. The potential for establishment in the North Sea region of all non-native species found was classified into three categories based on the degree of similarity of climatic conditions in the North Sea and the donor region. Based on this criterion 19 of the species found in the fouling communities on ships' hulls were deemed to have a high potential for establishment in the North Sea.     

Genetic variation, predator-prey interactions and food web structure

Food webs are networks of species that feed on each other. The role that within-population phenotypic and genetic variation plays in food web structure is largely unknown. Here, I show via simulation how variation in two key traits, growth rates and phenology, by influencing the variability of body sizes present through time, can potentially affect several structural parameters in the direction of enhancing food web persistence: increased connectance, decreased interaction strengths, increased variation among interaction strengths and increased degree of omnivory. I discuss other relevant traits whose variation could affect the structure of food webs, such as morphological and additional life-history traits, as well as animal personalities. Furthermore, trait variation could also contribute to the stability of food web modules through metacommunity dynamics. I propose future research to help establish a link between within-population variation and food web structure. If appropriately established, such a link could have important consequences for biological conservation, as it would imply that preserving (functional) genetic variation within populations could ensure the preservation of entire communities.     

Influence of pollution on the harpacticoid copepods of two North Sea estuaries

Seasonal monitoring of the meiobenthos in the Dutch estuaries revealed an anomaly in density and diversity of harpacticoid copepods in the Westerschelde.Another Dutch estuary, the Eems Dollard, has comparable hydrodynamical, physical and sedimentological, characteristics and a similar fauna, but even in the severely organically polluted oligohaline mudflats of this estuary, annual average density and diversity of endo-epibenthic harpacticoid communities are higher than at similar less enriched meso- to polyhaline biotopes of the Westerschelde.Besides the concentrations of inorganic pollutants, such as phosphates and nitrates, the concentrations of pesticides, cyanide, detergents, phenoles, oils, polychlorobenzenes, polycyclic aromatic hydrocarbons and heavy metals were compared in both estuaries and compared to suggested permissive levels when available. From this it appears that these pollutants are present in the Westerschelde either in too low concentrations to be considered dangerous or at concentrations comparable to those occurring in the Eems Dollard, except for heavy metals.The load of a.o. Zn, Cu and Pb is distinctly and persistently higher in sediments and suspensions of the Westerschelde than in the Eems Dollard and copper is continuously present in a concentration at which, according to bioassays, egg production and larval development of planktonic copepods are severely affected. The remarkable scarcity of harpacticoid life on nutrient rich mudflats of the Westerschelde is thus probably due to heavy metal pollution.Since no other hardbodied meio- and macrobenthic taxa nor the plankton of this estuary show such a marked impoverishment, benthic harpacticoids prove to be suitable as indicators for the first stages of ecosystem-breakdown in estuarine and coastal zones polluted by trace-metals.     

Seasonal variability of inorganic and organic nitrogen in the North Sea

This study considers the cycling of nitrogen in the waters of the North Sea, particularly focussing on organic nitrogen. Dissolved inorganic nitrogen (DIN), dissolved organic nitrogen (DON) and particulate organic nitrogen (PON) were measured in the North Sea over a one-year period (autumn 2004–summer 2005). The surface water concentrations of nitrate, ammonium, DON and PON during the present study ranged from <0.1–7.2 μM, <0.1–2.0 μM, 1.9–11.2 μM and 0.3–5.6 μM, respectively, with DON the dominant fraction of total nitrogen at all times. These nutrients concentrations were significantly lower compared to previous studies in the southern North Sea. The seasonal variations showed high mean surface concentrations of nitrate (4.7 ± 0.6 μM) and DON (8.9 ± 0.9 μM), low ammonium (<0.1 μM) and PON (0.8 ± 0.1 μM) in winter, shifting to low nitrate (0.3 ± 0.3 μM) and DON (4.2 ± 1.2 μM) in summer, with high ammonium (0.8 ± 0.4 μM) in autumn and PON (2.5 ± 1.2 μM) in spring. Highest mean surface DON concentration was measured in winter and may be due to resuspension of the organic matter from the bottom sediments. For autumn and spring, phytoplankton DON release was likely to be the most significant source of DON as shown by high concentrations of low molecular weight (LMW) DON and its positive correlation to chlorophyll a. Low total and LMW DON concentrations during summer were likely to be due to the uptake of the LMW DON fraction by phytoplankton and bacteria and the stratification of the water column. DON is therefore shown to be a potentially important source of nitrogen in shelf seas especially after the spring bloom has depleted nitrate to limiting concentrations. Handling editor: L. Naselli-Flores