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

The determination of trophic level for the biology in a marine ecosystem is very important as alteration of its structure and function may be reflected in the tro-phic level of component species. A change in trophic level indicates variation in an organism’s feeding bi-ology or in the pathway of energy flow from primary producers to the consumer. The gut content analysis is a traditional method for studying trophodynamics of food web in marine ecosystems. Species composition and amounts in al…     

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.     

Benthic food web structure in the southeastern Chukchi Sea: an assessment using ��13C and ��15N analyses

The benthos of the southeastern Chukchi Sea shelf is typified by high faunal abundance and biomass resulting from settlement of a large proportion of seasonal phytoplankton under highly nutritious offshore Bering Shelf Anadyr Water (BSAW). In contrast, inshore Alaska Coastal Water (ACW) is much less productive. Yet the Chukchi Bight and Kotzebue Sound, located under ACW in the southeastern Chukchi Sea, contain a substantial faunal abundance and biomass of invertebrates, fishes and marine mammals. We examined food web structure to gain an understanding of how a relatively rich benthic fauna with a high biomass can be supported under ACW with a supposedly low flux of carbon to the benthos. We measured stable isotope (δ¹³C and δ¹⁵N) values of selected organisms (from zooplankton to fishes) as markers of food sources and trophic position to compare fauna on the shelf under BSAW with that in the Chukchi Bight and Kotzebue Sound under ACW. Relative isotope position of organisms in all three regions was similar, even though some pelagic species within the Sound were depleted in δ¹³C compared to the other regions. We attribute the depletion to the influence of terrestrially derived carbon. We suggest that the hydrodynamics along an oceanic front between the Chukchi Shelf and the Chukchi Bight support the advection of nutrient-rich POC into the Bight and Sound as additional food sources to local production. We conclude that local conditions and multiple POC sources in the Bight and Sound support the substantial population of benthic invertebrates and the fishes, seabirds, and marine mammals that feed on them.     

Temporal and spatial changes in benthic invertebrate trophic networks along a taxonomic richness gradient

Species interactions underlie most ecosystem functions and are important for understanding ecosystem changes. Representing one type of species interaction, trophic networks were constructed from biodiversity monitoring data and known trophic links to assess how ecosystems have changed over time. The Baltic Sea is subject to many anthropogenic pressures, and low species diversity makes it an ideal candidate for determining how pressures change food webs. In this study, we used benthic monitoring data for 20 years (1980–1989 and 2010–2019) from the Swedish coast of the Baltic Sea and Skagerrak to investigate changes in benthic invertebrate trophic interactions. We constructed food webs and calculated fundamental food web metrics evaluating network horizontal and vertical diversity, as well as stability that were compared over space and time. Our results show that the west coast of Sweden (Skagerrak) suffered a reduction in benthic invertebrate biodiversity by 32% between the 1980s and 2010s, and that the number of links, generality of predators, and vulnerability of prey have been significantly reduced. The other basins (Bothnian Sea, Baltic Proper, and Bornholm Basin) do not show any significant changes in species richness or consistent significant trends in any food web metrics investigated, demonstrating resilience at a lower species diversity. The decreased complexity of the Skagerrak food webs indicates vulnerability to further perturbations and pressures should be limited as much as possible to ensure continued ecosystem functions.     

Contribution of sea ice microbial production to Antarctic benthic communities is driven by sea ice dynamics and composition of functional guilds

Organic matter produced by the sea ice microbial community (SIMCo) is an important link between sea ice dynamics and secondary production in near‐shore food webs of Antarctica. Sea ice conditions in McMurdo Sound were quantified from time series of MODIS satellite images for Sept. 1 through Feb. 28 of 2007–2015. A predictable sea ice persistence gradient along the length of the Sound and evidence for a distinct change in sea ice dynamics in 2011 were observed. We used stable isotope analysis (δ¹³C and δ¹⁵N) of SIMCo, suspended particulate organic matter (SPOM) and shallow water (10–20 m) macroinvertebrates to reveal patterns in trophic structure of, and incorporation of organic matter from SIMCo into, benthic communities at eight sites distributed along the sea ice persistence gradient. Mass‐balance analysis revealed distinct trophic architecture among communities and large fluxes of SIMCo into the near‐shore food web, with the estimates ranging from 2 to 84% of organic matter derived from SIMCo for individual species. Analysis of patterns in density, and biomass of macroinvertebrate communities among sites allowed us to model net incorporation of organic matter from SIMCo, in terms of biomass per unit area (g/m²), into benthic communities. Here, organic matter derived from SIMCo supported 39 to 71 per cent of total biomass. Furthermore, for six species, we observed declines in contribution of SIMCo between years with persistent sea ice (2008–2009) and years with extensive sea ice breakout (2012–2015). Our data demonstrate the vital role of SIMCo in ecosystem function in Antarctica and strong linkages between sea ice dynamics and near‐shore secondary productivity. These results have important implications for our understanding of how benthic communities will respond to changes in sea ice dynamics associated with climate change and highlight the important role of shallow water macroinvertebrate communities as sentinels of change for the Antarctic marine ecosystem.     

Trophic interactions within the Ross Sea continental shelf ecosystem

The continental shelf of the Ross Sea is one of the Antarctic's most intensively studied regions. We review the available data on the region's physical characteristics (currents and ice concentrations) and their spatial variations, as well as components of the neritic food web, including lower and middle levels (phytoplankton, zooplankton, krill, fishes), the upper trophic levels (seals, penguins, pelagic birds, whales) and benthic fauna. A hypothetical food web is presented. Biotic interactions, such as the role of Euphausia crystallorophias and Pleuragramma antarcticum as grazers of lower levels and food for higher trophic levels, are suggested as being critical. The neritic food web contrasts dramatically with others in the Antarctic that appear to be structured around the keystone species Euphausia superba. Similarly, we suggest that benthic-pelagic coupling is stronger in the Ross Sea than in most other Antarctic regions. We also highlight many of the unknowns within the food web, and discuss the impacts of a changing Ross Sea habitat on the ecosystem.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

北部湾生态通道模型的构建

根据1997年～1999年在北部湾进行的渔业资源和生态环境调查数据,利用EwE软件构建北部湾生态系统的营养通道模型,模型由16个功能组构成,包括了哺乳动物和海鸟,每一组都代表在生态系统中具有相似地位的有机体,基本覆盖了北部湾生态系统能量流动的主要过程.模型分析表明,北部湾生态系统的能量流动主要以捕食食物链途径为主,其中无脊椎动物在能量从低级向高层次转换中起关键作用.各功能组的营养级范围为1.00～4.04,哺乳动物占据了最高营养层.生态网络分析表明,系统的能量流动主要有6级,来自初级生产者的能流效率为12.2%,来自碎屑的转换效率为12.3%,平均能量转换效率为12.2%.模型估算的可利用的生物量密度为8.7 t·km^-2,生态系统的生物生产量只占系统净初级生产力的1.81%.当前北部湾海洋生态系统处于不稳定状态.     

Ecosystem functioning of two marine food webs in the North‐Western Ionian Sea (Central Mediterranean Sea)

The ecosystem functioning of two marine food webs covering the north‐eastern (Salento) and south‐western (Calabria) sectors of the North‐Western Ionian Sea (NWIS) (Central Mediterranean Sea) was investigated through a food‐web model. Data inputs covered a wide set of ecological information applied to 58 functional groups (FGs). The sum of consumption and the mean predation mortality rate were calculated for benthic, demersal, and pelagic subsystems indicating the predator and prey roles of the FGs. A complex system of energy and biomass exchanges characterized the investigated food webs indicating an important benthic‐pelagic coupling. In the food webs of both areas, the regulation of flows between the benthic‐pelagic coupling seems to occur through the benthopelagic shrimps and the small pelagics due to their wasp‐waist control role. Differences were observed concerning the top predators. Odontocetes play this keystone role in the Salento food web. Anglers, bathyal squids, and sharks assume this functional role in Calabria. The geomorphology and hydrography in the NWIS could affect the biomass and energy exchanges in this coupling. The higher flows of consumption of the benthic system observed in the Calabria food web could be influenced by a widespread presence of canyons along the continental edge which increase the benthic productivity. In contrast, the flows of consumption in the Salento food web seem to be driven by the planktonic productivity supporting the pelagic, benthopelagic, and demersal compartments. This condition could be favored by the large extension of the shelf break zone. The food‐web models realized for the NWIS represent ideal platforms for the development of analysis with dynamic simulations. The comparative analysis of the two food webs by means of the FGs and their functional traits allowed the general pattern of ecosystem structure and functioning in the NWIS to be identified, making it an interesting approach to investigate the marine ecosystem.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Southeastern Chukchi Sea (Alaska) epibenthos

Epibenthos of the southeastern Chukchi Sea, inclusive of Kotzebue Sound, was sampled in 1976. Crustaceans dominated abundance while echinoderms, mainly sea stars, dominated biomass. Spatial distribution of fauna was determined by cluster analysis. Scavenger-predators were dominant trophic groups, although suspension feeders dominated some regions. Unexpected high abundance and biomass under Alaska Coastal Water and factors related to presence and distribution of fauna are discussed. Dissimilarities between the benthic systems of the southeastern and northeastern Chukchi Seas are attributed to more complex water mass and flow patterns in the southern system. Entrained particulate organic carbon disperses through the area to support an abundant and apparently highly productive benthic fauna, which sustains resident and transient populations of demersal fishes and marine mammals. Epibenthos was sampled again in 1998 and, although little change was apparent in community composition, many taxa had higher abundance and biomass than 22 years earlier, a trend similar to findings observed in the northeastern Bering Sea.Electronic Supplementary Material Supplementary material is available for this article at     

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.     

Assessing the functional importance of large‐bodied invertebrates in experimental headwater streams

Recent theoretical advances in food web ecology emphasize the importance of body size disparities among species for the structure, stability and functions of ecosystems. Experimental confirmations of the functional importance of large species, independent of their trophic position, are scarce. We specifically examine the multiple ecological roles of large invertebrates from two distinct trophic levels in headwater streams. We experimentally manipulated the presence of large predatory invertebrates (two Perlid stoneflies) or detritivores (a limnephilid caddisfly and a Pteronarcys stonefly) in a two‐by‐two design in stream channels open to immigration/emigration of smaller biota. We assessed treatment effects on the trophic structure of the benthic invertebrate community, dynamics of basal resources (benthic algae and leaf litter of cedar and alder), and stability of litter decomposition rates against an experimental pulse perturbation (fine sediment input). The presence of the large invertebrates was associated with a ten‐fold decrease in the biomass of invertebrate filterers whereas other trophic groups were unaffected by the large species. The biomass of benthic algae was lower and the rate of mass loss of alder litter was higher in channels lacking the large predators, thus revealing trophic cascades operating along both algal‐based and detritus‐based food chains. The large predators had no detectable effect on the decomposition of cedar whereas both cedar and alder disappeared faster in the presence of the large detritivores. Furthermore, the large predators and large detritivores interactively influenced the decomposition of the cedar–alder mixture through a litter diversity effect and the variability of the rate of alder decomposition after a pulse of fine sediment. Because the large invertebrates affected multiple ecosystem properties, and as their absence was not rapidly compensated for by small immigrant species, our findings support the notion that large species could be critically important in controlling ecosystem structure and functioning.     

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).     

捕捞对北部湾海洋生态系统的影响

利用Ecopath with Ecosim (EwE) 5.1软件构建了北部湾海洋生态系统1959—1960年的Ecosim模型,包含渔业、海洋哺乳动物、海鸟、中上层鱼类、底层鱼类、底栖无脊椎动物等20个功能组,通过与1997—1999年调查数据对比,分析了捕捞活动对北部湾生态系统的结构和功能的影响.结果表明：近40年来在捕捞强度不断增加的压力下,生态系统的结构和功能发生显著变化,长寿命、高营养级的肉食性鱼类生物量下降明显,系统以短寿命、小型鱼类和无脊椎动物占优势.1999年的大中型鱼类的生物量仅为1960年的6%,而小型鱼类和无脊椎动物则明显上升,尤其是头足类生物量上升了2.7倍,渔获物的营养级则从1960年的3.2降低到1999年的298,体现了“捕捞降低海洋食物网”的特点,目前的开发模式是不可持续的.利用20世纪90年代数据预测了降低捕捞压力后生态系统的变化.本研究证实了可以使用Ecosim模型预测捕捞压力对生态系统的影响.     

Macroecological patterns and niche structure in a new marine food web

The integration of detailed information on feeding interactions with measures of abundance and body mass of individuals provides a powerful platform for understanding ecosystem organisation. Metabolism and, by proxy, body mass constrain the flux, turnover and storage of energy and biomass in food webs. Here, we present the first food web data for Lough Hyne, a species rich Irish Sea Lough. Through the application of individual-and size-based analysis of the abundance-body mass relationship, we tested predictions derived from the metabolic theory of ecology. We found that individual body mass constrained the flux of biomass and determined its distribution within the food web. Body mass was also an important determinant of diet width and niche overlap, and predator diets were nested hierarchically, such that diet width increased with body mass. We applied a novel measure of predator-prey biomass flux which revealed that most interactions in Lough Hyne were weak, whereas only a few were strong. Further, the patterning of interaction strength between prey sharing a common predator revealed that strong interactions were nearly always coupled with weak interactions. Our findings illustrate that important insights into the organisation, structure and stability of ecosystems can be achieved through the theoretical exploration of detailed empirical data.     

Seasonal microbial processes in a high-latitude fjord (Kongsfjorden, Svalbard): I. Heterotrophic bacteria, picoplankton and nanoflagellates

Temporal dynamics of the microbial food web in the Barents Sea and adjacent water masses in the European Arctic are to a large extent unknown. Seasonal variation in stocks and production rates of heterotrophic bacteria and phototrophic and heterotrophic picoplankton and nanoflagellates was investigated in the upper 50 m of the high-latitude Kongsfjorden, Svalbard, during six field campaigns between March and December 2006. Heterotrophic bacteria, picoplankton and nanoflagellates contributed to ecosystem structure and function in all seasons. Activity within the microbial food web peaked during spring bloom in April, parallel to low abundances of mesozooplankton. In the nutrient-limited post-bloom scenario, an efficient microbial loop, fuelled by dissolved organic carbon from abundant mesozooplankton feeding on phytoplankton and protozooplankton, facilitated maximum integrated primary production rates. A tight microbial food web consisting of heterotrophic bacteria and phototrophic and heterotrophic picoplankton and nanoflagellates was found in the stratified water masses encountered in July and September. Microbial stocks and rates were low but persistent under winter conditions. Seasonal comparisons of microbial biomass and production revealed that structure and function of the microbial food web were fundamentally different during the spring bloom when compared with other seasons. While the microbial food web was in a regenerative mode most of the time, during the spring bloom, a microbial transfer mode represented a trophic link for organic carbon in time and space. The microbial food web’s ability to fill different functional roles in periods dominated by new and regenerated production may enhance the ecological flexibility of pelagic ecosystems in the present era of climate change.     

Human activities as a driver of spatial variation in the trophic structure of fish communities on Pacific coral reefs

Anthropogenic activities such as land‐use change, pollution and fishing impact the trophic structure of coral reef fishes, which can influence ecosystem health and function. Although these impacts may be ubiquitous, they are not consistent across the tropical Pacific Ocean. Using an extensive database of fish biomass sampled using underwater visual transects on coral reefs, we modelled the impact of human activities on food webs at Pacific‐wide and regional (1,000s–10,000s km) scales. We found significantly lower biomass of sharks and carnivores, where there were higher densities of human populations (hereafter referred to as human activity); however, these patterns were not spatially consistent as there were significant differences in the trophic structures of fishes among biogeographic regions. Additionally, we found significant changes in the benthic structure of reef environments, notably a decline in coral cover where there was more human activity. Direct human impacts were the strongest in the upper part of the food web, where we found that in a majority of the Pacific, the biomass of reef sharks and carnivores were significantly and negatively associated with human activity. Finally, although human‐induced stressors varied in strength and significance throughout the coral reef food web across the Pacific, socioeconomic variables explained more variation in reef fish trophic structure than habitat variables in a majority of the biogeographic regions. Notably, economic development (measured as GDP per capita) did not guarantee healthy reef ecosystems (high coral cover and greater fish biomass). Our results indicate that human activities are significantly shaping patterns of trophic structure of reef fishes in a spatially nonuniform manner across the Pacific Ocean, by altering processes that organize communities in both “top‐down” (fishing of predators) and “bottom‐up” (degradation of benthic communities) contexts.     

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter‐specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic‐feeding ice‐associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983–1999 and 2000–2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic‐level productivity in these marine ecosystems.