Trophic network model of the Northern Adriatic Sea: Analysis of an exploited and eutrophic ecosystem

A quantitative model of the trophic network of Northern Adriatic Sea marine ecosystem during the 1990s has been constructed, with the goal of analysing its trophic structure, identifying the key trophic groups and assessing the anthropogenic impacts on the ecosystem using the Ecopath modelling protocol. The Northern Adriatic Sea is an eutrophic, shallow basin, and one of the most heavily fished areas in the Mediterranean Sea. The network aggregation into discrete trophic levels sensu Lindeman shows that low trophic levels dominate biomass and energy flows, with 40% of the total system throughput flowing out from trophic level 2. Instead, upper trophic levels appear bottom-up controlled, highly depleted and not exerting any control on the trophic network, as shown by mixed trophic impact-based analyses. Microbial loop is comparable to grazing with respect to the magnitude of flows involved, as 66% of the trophic network flows originate from detritus, which is mainly consumed by bacteria. Key trophic groups are plankton groups, macro-crustaceans and detritus, and other r-selected organisms like squids and small pelagics, which have a great influence on the ecosystem. In particular, zooplankton acts as a bottleneck for energy flows, limiting the energy from the low trophic levels effectively reaching the upper food web. The high pelagic production caused by eutrophication sustains high fishery landings and impressive discard quantities, as well as the benthic compartment. Overall, the ecosystem appears quite productive and in a stressed and developmental status. Model results and comparisons with few existing historical data suggest that the low maturity and stressed state of the Northern Adriatic Sea are not only due to natural characteristics, but mainly to anthropogenic pressures.     

中街山列岛海域食物网结构初步研究及主要消费者潜在碳源分析

为了解中街山列岛的食物网结构特征,本研究于2020年7月在中街山列岛海域采集鱼类、虾蟹类、头足类、贝螺类和浮游动物等消费者样本,利用碳、氮稳定同位素技术,分析大型海藻、浮游植物、悬浮颗粒有机物(POM)和沉积物有机物(SOM)4种潜在碳源对消费者的贡献率以及中街山列岛海域的食物网结构和营养关系。结果表明：(1)碳源的δ¹³C值范围在-22.93‰～-9.73‰之间,δ15N值范围在1.72‰～7.68‰之间,消费者的δ¹³C值范围在-21.95‰～-12.55‰之间,δ15N值范围在4.13‰～12.92‰之间,不同碳源及不同消费者类群之间的碳、氮稳定同位素均有显著性差异(p<0.01);(2)应用SIBER模型计算中街山列岛海域的营养结构指标,与其他海域的研究结果对比发现,该海域生态系统的食源多样性更加丰富,营养级长度和生态位总空间较高;(3)应用SIAR模型计算碳源贡献率,结果表明浮游植物和POM是该海域的重要碳源,平均贡献率为29.63%和28.72%;浮游植物对浮游动物的贡献率最大为80.58%,POM对鱼类的贡献率最大为79....     

Trophic structure and functioning in a eutrophic and poorly flushed lagoon in southwestern Taiwan

Tapong Bay, a eutrophic and poorly flushed tropical lagoon, supports intensive oyster culture. Using the Ecopath approach and network analysis, a mass-balanced trophic model was constructed to analyze the structure and matter flows within the food web. The lagoon model is comprised of 18 compartments with the highest trophic level of 3.2 for piscivorous fish. The high pedigree index (0.82) reveals the model to be of high quality. The most-prominent living compartment in terms of matter flow and biomass in the lagoon is cultured oysters and bivalves, respectively. The mixed trophic impacts indicate that phytoplankton and periphyton are the most-influential living compartments in the lagoon. Comparative analyses with the eutrophic and well-flushed Chiku Lagoon and non-eutrophic tropical lagoons show that high nutrient loadings might stimulate the growth and accumulation of phytoplankton and periphyton and therefore support high fishery yields. However, net primary production, total biomass, fishery yields per unit area, and mean transfer efficiency of Tapong Bay were remarkably lower than those of Chiku Lagoon. The lower transfer efficiency likely results from the low mortality of cultured oysters and invasive bivalves from predation or the lower density of benthic feeders constrained by the hypoxic bottom water as a result of poor flushing. This might therefore result in a great proportion of flows to detritus. However, the hypoxic bottom water might further reduce the recycling of the entering detritus back into the food web. In contrast to many estuaries and tropical lagoons, poor flushing of this eutrophic tropical lagoon might induce a shift from detritivory to herbivory in the food web.     

Quantitative model of trophic interactions in Beibu Gulf ecosystem in the northern South China Sea

1IntroductionThe Beibu Gulf is a natural semiclosed conti-nental sea of the South China Sea,which is situatedat17°00′~21°45′N,105°40′~110°10′E,and sur-rounded by China and Vietnam(see Fig.1).It hasa subtropic monsoon climate with an average winter     

Redundancy and diversity of functional reef fish groups of the Mexican Eastern Pacific

Coral reefs are in crisis worldwide. Fish play a key role in the resilience of reefs because most of the biomass of the ecosystem flows through them. Evidence indicates that the response to stressors will vary markedly depending on the resilience of the ecosystem. In order to assess the functional groups of reef fishes, morphometric and ecomorphological analyses were carried out on the shape and diet of 117 reef fish species from the Mexican Eastern Pacific (MEP) from 23°N to 15°N, spanning the years 2009–2012. Six trophic groups were identified and subdivided into 19 functional groups: (A) detritus and plants consumers, three morphologies; (B) plants and zoobenthos consumers, four morphologies; (C) zoobenthos consumers, two morphologies; (D) zoobenthos and zooplankton consumers, three morphologies; (E) zooplankton and nekton consumers, three morphologies; and (F) nekton consumers, four morphologies. Our main conclusions are: (i) fish communities of the MEP have a high degree of ecological redundancy; (ii) fish diversity is not related to the coral cover; and (iii) the combination of using both morphology and diet produce a more effective classification of functional groups. Future studies such as these will allow a deeper understanding of environment and it will help illuminate the effects of fish morphology and diet on population structure.     

Changes in trophic flows and ecosystem properties of the Beibu Gulf ecosystem before and after the collapse of fish stocks

Mass-balance models (Ecopath) of the ecosystem before and after collapse (1959-1961 and 1997-1999) of fish stocks were developed with Ecopath software to compare the differences in ecosystem structure, functioning and ecosystem properties of the Beibu Gulf. The model includes 20 functional groups consisting of commercial important fish groups and other ecologically important groups in the ecosystem such as zooplankton, phytoplankton, and detritus. Results indicated that biomass and catches of the system have changed drastically between the 1960s and 1990s, especially for the high trophic levels (TL). The biomass of level V in the early 1960s was 32 times higher than that of the late 1990s, however, the biomass of level I and II in the 1990s was higher than the 1960s. Despite the higher catches in the 1990s, fishing was ecologically less expensive during the 1990s than 1960s due to small fish catches were large. Mean transfer efficiency decreased from for 10.2% in the 1960s to 9.1% in the 1990s periods. According to the summary statistics, the parameters of net system production (NPS) and total primary production to total respiration ratio were increased from 1.013 in the 1960s to 2.184 in the 1990s, however, the connectance index (CI), system omnivore index, Finn’s cycling index and mean path length decreased from the 1960s to the 1990s. The overhead (O) was higher in the 1990s model while the ascendancy (A) decreased nearly 10% in the 1960s. The ‘Keystoneness’ result indicate that zooplankton was identified as keystone species in 1960s, however, the elasmobranches was keystone species in the late 1990s. The average trophic level of the fishery decreased from 3.32 in the 1960s to 2.98 in the 1990s, and exhibits classic symptoms of “fishing down the food web”. All the indices of the system attributes suggests that the Beibu Gulf ecosystem in 1960s was found to be more mature than in the 1990s due to the collapse of demersal ecosystem, and the ecosystem changed from being dominated by long-lived, high trophic level groundfish dominated system toward a system with small-size and low-value species over fifty years.     

Management of the coastal-sea-zone-ecosystem model

We present a model of the processes of development in the ecosystem of the coastal zone of the sea constructed by the method of adaptive balance of causes (ABC-method). The model contains nine variables (phyto- and zooplankton, fish larvae, fishes, oxygen, carbon dioxide, salinity, biogenic elements, and detritus) affected by the solar radiation, surface wind, sea temperature, and river discharge. The equations of the model contain control agents governing the dependences of the elements of the food chain on the vitally important resources and environmental conditions. The scenarios of the processes of development are constructed for the simulated annual courses of external actions. The possibility of management of fish resources by regulating the flows of biogenic elements and detritus with river discharge is discussed. A mechanism of management based on the monitoring of the concentration of dissolved oxygen in seawater is proposed. The numerical experiments show that the ABC-model of the processes of development in the ecosystem of the coastal zone of the sea makes it possible to simulate the scenarios of regulation of the flows of biogenic elements and detritus into the sea with an aim of preservation of of fish resources. Translated from Morskoi Gidrofizicheskii Zhurnal, No. 1, pp. 36–49, January–February, 2009.     

Trophic modeling of the Northern Humboldt Current Ecosystem, Part I: Comparing trophic linkages under La Niña and El Niño conditions

The El Niño of 1997–98 was one of the strongest warming events of the past century; among many other effects, it impacted phytoplankton along the Peruvian coast by changing species composition and reducing biomass. While responses of the main fish resources to this natural perturbation are relatively well known, understanding the ecosystem response as a whole requires an ecotrophic multispecies approach. In this work, we construct trophic models of the Northern Humboldt Current Ecosystem (NHCE) and compare the La Niña (LN) years in 1995–96 with the El Niño (EN) years in 1997–98. The model area extends from 4°S–16°S and to 60 nm from the coast. The model consists of 32 functional groups of organisms and differs from previous trophic models of the Peruvian system through: (i) division of plankton into size classes to account for EN-associated changes and feeding preferences of small pelagic fish, (ii) increased division of demersal groups and separation of life history stages of hake, (iii) inclusion of mesopelagic fish, and (iv) incorporation of the jumbo squid (Dosidicus gigas), which became abundant following EN. Results show that EN reduced the size and organization of energy flows of the NHCE, but the overall functioning (proportion of energy flows used for respiration, consumption by predators, detritus and export) of the ecosystem was maintained. The reduction of diatom biomass during EN forced omnivorous planktivorous fish to switch to a more zooplankton-dominated diet, raising their trophic level. Consequently, in the EN model the trophic level increased for several predatory groups (mackerel, other large pelagics, sea birds, pinnipeds) and for fishery catch. A high modeled biomass of macrozooplankton was needed to balance the consumption by planktivores, especially during EN condition when observed diatoms biomass diminished dramatically. Despite overall lower planktivorous fish catches, the higher primary production required-to-catch ratio implied a stronger ecological impact of the fishery and stresses the need for precautionary management of fisheries during and after EN. During EN energetic indicators such as the lower primary production/total biomass ratio suggest a more energetically efficient ecosystem, while reduced network indicators such as the cycling index and relative ascendency indicate of a less organized state of the ecosystem. Compared to previous trophic models of the NHCE we observed: (i) a shrinking of ecosystem size in term of energy flows, (ii) slight changes in overall functioning (proportion of energy flows used for respiration, consumption by predators and detritus), and (iii) the use of alternate pathways leading to a higher ecological impact of the fishery for planktivorous fish.     

Sources and transfer of organic matter in food webs of a Mediterranean coastal environment: Evidence for spatial variability

The spatial variability in the food web structure of a Mediterranean semi-enclosed coastal environment (Stagnone di Marsala, Italy) was investigated using stable carbon and nitrogen isotopes. Organic matter sources and consumers were sampled in two locations with different environmental features (e.g. hydrodynamic regime, open-sea influence, vegetal coverage). Overall more ¹³C-enriched and ¹⁵N-depleted values were found in the central location than in the southern for organic matter sources and consumers. Pelagic consumers (zooplankton and juveniles of transient fish) showed slight spatial differences and in both locations seemed to depend on phytoplankton as the ultimate energy source. In contrast, benthic consumers (epifauna and resident fish) exhibited remarkable differences between locations. Spatial differences in organic matter sources were smaller than in benthic consumers and thus consumers presumably exploited different ultimate organic matter sources in the two locations. Sedimentary organic matter and epiphytes appeared to be the main primary producers transferred within the food web in both locations, and seagrasses seemed to play a non-negligible trophic role in the central location. The results of this paper corroborate the finding food webs are characterised by high spatial variability even on a small spatial scale and environmental heterogeneity more than primary production that seems to influence the trophic role of autotrophs.     

应用物质平衡模型模拟一个半封闭海湾的营养结构和能流特征

The marine ecosystem of the Jiaozhou Bay has degraded significantly in fisheries productivity and its ecological roles as spawning and nursery ground for many species of commercial importance has been declining in recent years. A mass-balanced trophic model was developed using Ecopath with Ecosim to evaluate the trophic structure of the Jiaozhou Bay for improving ecosystem management. The model were parameterized based on the fisheries survey data in the Jiaozhou Bay in 2011, including 23 species groups and one detritus group according to their ecological roles. The trophic levels of these ecological groups ranged from 1(primary producers and detritus) to4.3(large demersal fishes). The estimated total system throughput was 12 917.10 t/(km~2·a), with 74.59% and25.41% contribution of the total energy flows from phytoplankton and detritus, respectively. Network analyses showed that the overall transfer efficiency of the ecosystem was 14.4%, and the mean transfer efficiency was 14.5%for grazing food chain and 13.9% for detritus food chain. The system omnivory index(SOI), Finn's cycled index(FCI) and connectance index(CI) were relatively low in this area while the total primary production/total respiration(TPP/TR) was high, indicating an immature and unstable status of the Jiaozhou Bay ecosystem. Mixed trophic impact analysis revealed that the cultured shellfish had substantial negative impacts on most functional groups. This study contributed to ecosystem-level evaluation and management planning of the Jiaozhou Bay ecosystem.     

印太交汇区浮游植物和浮游动物生态学研究进展

海洋浮游植物和浮游动物是海洋生态系统中的重要组成部分,支撑了整个海洋生态系统的正常运转.因此,海洋浮游植、动物的生态学研究有利于我们全面认识和了解一个海洋生态系统的状况.印太交汇区作为全球最大的海洋生物多样性中心,是国际上生物多样性研究的热点区域,但该区域对浮游生物生态学方面的研究较少,不利于我们深入认识该区域生物多样...     

A Preliminary Trophic Model of the Continental Shelf, South-western Gulf of Mexico

A preliminary mass-balance trophic model was constructed to determine the flow of energy in a community of fish and invertebrates on the continental shelf of the south-western Gulf of Mexico. Input parameters were taken from the literature, except for the biomass of fish groups which was obtained from trawl surveys in the study area. The model consists of 12 fish groups, five invertebrate groups, phytoplankton and detritus. Results indicate an imbalance between primary production and consumption, with only about 10% of primary production being consumed in the water column. Most of the primary production is exported to detritus which forms the basis of the food-web, with a detritivory/herbivory ratio of 2·5. Benthic invertebrates play a significant role in transferring energy from detritus to higher trophic levels. Eight discrete trophic levels were found, with very reduced flow at levels higher than the fifth. The highest fractional trophic level is 4·3, and consists of sharks. Analysis of mixed trophic impacts showed that detritus and lower trophic levels have a significant positive impact on other groups in the system. Mean transfer efficiency is 9·2%. Some whole system properties are also given. Of net primary production, 6·7% is required to sustain current catch levels, suggesting that the resources in this area are fully exploited.     

A nitrogen and phosphorus dynamic model of mesocosm pelagic ecosystem in the Jiaozhou Bay in China

A nitrogen and phosphorus dynamic model of mesocosm pelagic ecosystem was established according to the summary and synthesis of the models available, in which seven state variables （DIN, PO4-P, DON, DOP, phytoplankton, zooplankton and detritus） were included. Logically it had five modules--phytoplankton, zooplankton, dissolved inorganic nutrients, dissolved organic nutrients and detritus. The results showed that this model could simulate the variations of DIN, PO4-P, DON, DOP, POC and phytoplankton biomass in pelagic ecosystem in mesocosm properly, based on the site experiment data in the Jiaozhou Bay in the autumn of 1999 and the summer of 2000. Not only the logical structure but also the model parameters were feasible, and about 20 parameters were made to fit for the Jiaozhou Bay during the simulation. All of these are necessary to study the control mechanism of nutrients biogeochemical cycling in the Jiaozhou Bay and other China＇ s coastal waters.     

Grazing experiments and model simulations of the role of zooplankton in Phaeocystis food webs

A combined empirical and modelling study was conducted to further examine the potential importance of grazing by zooplankton in pelagic food webs in which Phaeocystis is a significant or dominant component. Laboratory experiments were designed to measure ingestion of Phaeocystis and other potential prey items which co-occur with Phaeocystis. Grazers included copepods and ciliates, and prey included Phaeocystis colonies and solitary cells, diatoms, ciliates, bacteria, and detritus. These data were expressed in the model currency of nitrogen units, and fit to hyperbolic tangent equations which included minimum prey thresholds. These equations and literature data were used to constrain a food web model whose purpose was to investigate trophic interactions rather than to mimic actual events. Nevertheless, the model output was similar to the general pattern and magnitude of development of Phaeocystis–diatom communities in some environments where they occur, e.g. north Norwegian waters. The model included three forms of nitrogen, three phytoplankton groups, bacteria, two zooplankton groups, and detritus, with detailed flows between compartments. An important component of the model was inclusion of variable prey preferences for zooplankton. The experiments and model simulations suggest several salient conclusions. Phaeocystis globosa colonies were eaten by a medium-sized copepod species, but ingestion appeared to be strongly dependent upon a proper size match between grazer and prey. If not, colonies were eaten little if at all. Phaeocystis solitary cells were ingested rapidly by ciliate microzooplankton, in agreement with prior literature observations. In contrast, detritus was eaten comparatively slowly by both ciliates and copepods. Both types of zooplankton exhibited apparent minimum prey thresholds below which grazing did not occur or was inconsequential. Model simulations implied that transitions between life cycle stages of Phaeocystis may potentially be important to phytoplankton–zooplankton interactions, and that relative rates of ingestion of Phaeocystis by various zooplankton may have significant impacts upon material fluxes through and out of Phaeocystis–diatom ecosystems. Indirect effects of trophic interactions appear to be equally significant as direct effects.     

福州沿海龟足的食性研究

龟足（Capitulum mitella Linnaeus）的食物组成可分为4类,动物性食物、植物性食物、有机碎屑和无机颗粒。动物性食物主要以桡足类为主,植物性食物主要以硅藻类为主,有机碎屑在龟足的食物组成中占据重要地位。龟足食物的食物多样性指数在夏季最高,冬季最低。小规格龟足主要摄食植物性食物和有机碎屑,未见摄食动物...     

Food web structure of the coastal area adjacent to the Tagus estuary revealed by stable isotope analysis

The identification of energy sources, pathways and trophic linkages among organisms is crucial for the understanding of food web dynamics. Stable isotopes were used to identify the trophic level of food web components and track the incorporation of organic matter of different origins in the coastal ecosystem adjacent to the Tagus estuary. It was shown that the river Tagus is a major source of organic carbon to this system. Also, the wide difference in δ¹³C among the primary consumers allowed the identification of the pelagic and the benthic energy pathways. The maximum trophic level observed was 2.4 for Sepia officinalis. This value is indicative of a short food web. It was concluded that the diet of the upper trophic level species relies directly on the lower food web levels to a considerable extent, instead of relying mostly on intermediate trophic level species. Moreover, the δ¹⁵N values of primary consumers were very close to that of particulate organic matter, probably due to poorly known processes occurring at the basis of the food web. This lowers the trophic length of the whole food web. Reliance on benthic affinity prey was high for all upper trophic level secondary consumers.     

Numerical modelling of spatio-temporal variability of growth of Mytilus edulis (L.) and influence of its cultivation on ecosystem functioning

One of the key needs of the aquaculture industry is the implementation of effective management methods to ensure the sustainability, economic viability and minimization of negative impacts on both human and ecosystem well-being. The authors developed a Fortran 90 implementation of the dynamic energy budget (DEB) model for Mytilus edulis. The model has been further developed to include physiological interactions with the ecosystem and coupled to a biogeochemical nutrient–phytoplankton–zooplankton–detritus (NPZD) model. Phytoplankton and detritus uptakes, oxygen utilisation, CO₂ production, NH₄ excretion, egestion of faeces, and assimilation of food are modelled. A novel approach was derived that accounts for the allocation of C and N in mussel flesh and shell organic fraction. The DEB–NPZD model has been subsequently coupled to a high resolution three dimensional numerical coastal ocean model of the south–west coast of Ireland, where approximately 80% of national rope mussel is produced annually. Simulations have been carried out for the time period July 2010–June 2011, for which the field data on mussel biometrics and ambient seawater properties were collated. The model accurately reproduced the spatio-temporal variability in blue mussel growth. It is also shown that the ecosystem dynamics is affected by the presence of aquaculture farms. The modelling system presented allows for the assessment of the impacts of aquaculture activities on water quality, quantification of the production and ecological carrying capacities and improvement of our understanding of the ecosystem functioning with particular emphasis on interactions between various trophic levels.     

A trophic model of the Pearl River Delta coastal ecosystem

A 26-compartment steady-state trophic model (1997–1999) was constructed using the Ecopath with Ecosim software to study the general status and development trends of the Pearl River Delta coastal ecosystem. The results show that the values of effective trophic level ranged from 1.00 to 4.21. It was found that a high trophic niche overlap existed in the typical estuarine ecosystem. Mixed trophic impacts show that detritus and the groups at the low trophic levels had positive influences on most groups. The ecosystem was found to be in an immature state during 1997–1999 based on the system statistics.     

Exergy,ecosystem functioning and efficiency in a coastal lagoon: The role of auxiliary energy

Exergy, as the sum of energy and information contained in a given system due to living organisms, can act as a quality indicator of ecosystems. Here, we investigated the exergy of Marsala Lagoon (Mediterranean Sea), along with microbial (prokaryotic and heterotrophic nanobenthos) biomass, prokaryotic heterotrophic production and extracellular enzymatic activities, and the biochemical composition of sediment organic matter. The aim of the study was to assess the role of auxiliary energy (e.g. hydrodynamic stress) in the ecosystem functioning and efficiency of a ‘detritus sink’ lagoon. Samples were collected at sites characterized by contrasting hydrodynamic and trophic conditions. Exergy transfer through the benthic microbial loop was influenced by two main factors: (1) organic matter bioavailability; and (2) hydrodynamic forcing. At both sites, the values of total exergy were higher in summer than in winter, and the specific exergy decreased from winter to autumn, along with increasing auxiliary energy. Our data indicate that in coastal ‘detritus sink’ systems, auxiliary energy sources can have a crucial role in exergy transfer and ecosystem functioning through modifying the efficiency of transfer to higher trophic levels of the refractory organic detritus, which is otherwise lost by burial in the sediment. As coastal lagoons are often intensively modified by human activities, we conclude that maintenance of the natural hydrodynamic regimes is a key factor in the preservation of the functioning of lagoon ecosystems and of their provision of goods and services to humans.     

Sensitivity analysis of a coastal marine ecosystem

An ecosystem with four compartments, i.e. PO₄-P, phytoplankton, zooplankton and detritus, in a coastal region was considered. Sensitivity analysis of a phosphorus flow model in well-mixed water without currents concludes that (1) the maximal photosynthetic rate,V _m , in Michaelis-Menten relation plays an important role in the distribution of biomass among compartments but the half saturation constant,K _s , is not so important, (2) the natural death rate of phytoplankton is important for the ecosystem, (3) the natural death rate of zooplankton is also effective on the ecosystem.A numerical experiment was also performed on the ecosystem with four compartments are also studied using a dynamical barotropic model of tidal currents of Mikawa Bay (Japan). Diffusion coefficient in diffusion equation plays the role of a linear smoothing parameter in the horizontal distribution of compartment. On the other hand, perturbations of biological parameters cause nonlinear variations in the horizontal distribution of compartment.