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.     

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.     

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.     

热带典型珊瑚岛礁海洋牧场花刺参底播增殖容量及其生态效应预测

为改善热带珊瑚岛礁型海洋牧场的珊瑚礁生境,实现生物资源的养护和渔业资源的产出功能,在对海参等高值经济种开展底播增殖前,科学评估其生态容量是防止引发海洋牧场生态风险的重要保证。运用生态系统模型法评估了三亚蜈支洲岛热带珊瑚岛礁海洋牧场花刺参(Stichopus monotuberculatus)的底播增殖容量。根据2020~2021年蜈支洲岛海洋牧场近岛区渔业资源调查与环境因子数据,运用Ecopath with Ecosim 6.6软件构建了该海域的生态系统营养通道模型。研究表明:生态系统各功能组营养级范围介于1~3.52,系统的食物网结构以牧食食物链为主,总能流中有43%的能量来源于碎屑功能组,其在系统总能流中有重要地位。系统的总平均能量传递效率为9.353%,略低于林德曼能量传递效率(10%)。总初级生产量/总呼吸量为3.726,总初级生产量/总生物量为28.834,系统连接指数为0.256,杂食性指数为0.120,系统Finn''s循环指数和平均路径长度分别为2.485%和2.379,表明近岛区生态系统食物网结构较为简单,且系统稳定性和成熟度偏低,易受外界干扰。根据模型评估的花刺参增殖生态容量为110.21 t/km²,是现存量的206 倍,有较大增殖空间,并且达到生态容量后碎屑组的能量再循环利用效率将显著增加,营养级结构能得到进一步优化,系统稳定性及成熟度将有所提高。基于研究结果,可适当采捕与花刺参生态位相近的生物,同时增殖放流其他处于不同营养层次的经济种,从而减少种间竞争,有效利用系统冗余能量,进而扩大花刺参的生态容量,实现海洋牧场的健康可持续发展。     

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.     

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.     

海洋自然资本评估中生物能值转换率的研究

自然资本评估是研究生态系统服务与社会经济发展的重要方式。由于海洋具有特殊性和复杂性、开放性、流动性和多层次耦合性等特点,海洋自然资本评估方法的研究进展缓慢。能值分析理论通过能值转换率将生态系统内流动和储存的各种不同类别的能量和物质转换为同一标准的能值,可衡量和比较不同等级的能量价值,系统完整地反映自然资本价值,这一特点是其他评估方法所缺乏的。为精确量化海洋生物资本价值,本研究引入海洋食物网信息能流图和生态系统能量传递规律,提出了生物能值转换率的经验公式,即Tn=T1·E^1-nL(T1为初级生产者的太阳能值转换率,n为营养级,EL为林德曼效率)。为验证经验公式的可行性,选用相关案例进行了研究,得到海洋食物网中不同食性生物所处营养级及其相应的能值转换率。经验公式有一定误差,具体应用时应通过调研文献和相关数据,确定关键参数的取值范围以提高准确性。本研究得出的经验公式简化了海洋生物能值转换率的计算方法,促进了能值分析理论在海洋自然资本评估中的进一步应用。     

Trophic structure and flows of energy in the Huizache–Caimanero lagoon complex on the Pacific coast of Mexico

The Huizache–Caimanero coastal lagoon complex on the Pacific coast of Mexico supports an important shrimp fishery and is one of the most productive systems in catch per unit area of this resource. Four other less important fish groups are also exploited. In this study, we integrated the available information of the system into a mass-balance trophic model to describe the ecosystem structure and flows of energy using the E approach. The model includes 26 functional groups consisting of 15 fish groups, seven invertebrate groups, macrophytes, phytoplankton, and a detritus group. The resulting model was consistent as indicated by the output parameters. According to the overall pedigree index (0.75), which measures the quality of the input data on a scale from 0 to 1, it is a high quality model. Results indicate that zooplankton, microcrustaceans, and polychaetes are the principal link between trophic level (TL) one (primary producers and detritus) and consumers of higher TLs. Most production from macrophytes flows to detritus, and phytoplankton production is incorporated into the food web by zooplankton. Half of the flow from TL one to the next level come from detritus, which is an important energy source not only for several groups in the ecosystem but also for fisheries, as shown by mixed trophic impacts. The Huizache–Caimanero complex has the typical structure of tropical coastal lagoons and estuaries. The TL of consumers ranges from 2.0 to 3.6 because most groups are composed of juveniles, which use the lagoons as a nursery or protection area. Most energy flows were found in the lower part of the trophic web.     

基于Ecopath模型的舟山海域褐菖鲉(Sebastiscus marmoratus)生态容量评估

根据2021年渔业资源调查数据构建了含有23个功能组的舟山海域生态系统Ecopath模型,分析了当前舟山海域生态系统总体特征并估算了褐菖鲉在舟山海域的生态容量。结果表明:舟山海域生态系统营养级范围为1.000 (浮游植物和有机碎屑)～4.277 ( 鳐类),石首鱼科、虾类和 鳐类为舟山海域生态系统中的关键种。碎屑食物链和牧食食物链是舟山海域生态系统主要的食物链。碎屑和浮游植物对食物网的贡献率分别为61.32%和38.69%。始于浮游植物和碎屑的营养传递效率分别是9.34%和10.50%,系统总营养传递效率是9.82%。总初级生产量/总呼吸量为2.26,系统连接指数为0.372,系统杂食性指数为0.222。生态系统总体特征反映了舟山海域生态系统的成熟状态较低,生态系统处于不稳定阶段,容易受到外界环境变化的影响。根据模型估算,当褐菖鲉生物量增加至8.6倍时,褐菖鲉达到生态容量0.007 95 t/km²,此时生态系统仍保持平衡,且生态系统总体特征基本稳定。因此,褐菖鲉在舟山海域尚有较大增殖潜力。     

Towards end-to-end models for investigating the effects of climate and fishing in marine ecosystems

End-to-end models that represent ecosystem components from primary producers to top predators, linked through trophic interactions and affected by the abiotic environment, are expected to provide valuable tools for assessing the effects of climate change and fishing on ecosystem dynamics. Here, we review the main process-based approaches used for marine ecosystem modelling, focusing on the extent of the food web modelled, the forcing factors considered, the trophic processes represented, as well as the potential use and further development of the models. We consider models of a subset of the food web, models which represent the first attempts to couple low and high trophic levels, integrated models of the whole ecosystem, and size spectrum models. Comparisons within and among these groups of models highlight the preferential use of functional groups at low trophic levels and species at higher trophic levels and the different ways in which the models account for abiotic processes. The model comparisons also highlight the importance of choosing an appropriate spatial dimension for representing organism dynamics. Many of the reviewed models could be extended by adding components and by ensuring that the full life cycles of species components are represented, but end-to-end models should provide full coverage of ecosystem components, the integration of physical and biological processes at different scales and two-way interactions between ecosystem components. We suggest that this is best achieved by coupling models, but there are very few existing cases where the coupling supports true two-way interaction. The advantages of coupling models are that the extent of discretization and representation can be targeted to the part of the food web being considered, making their development time- and cost-effective. Processes such as predation can be coupled to allow the propagation of forcing factors effects up and down the food web. However, there needs to be a stronger focus on enabling two-way interaction, carefully selecting the key functional groups and species, reconciling different time and space scales and the methods of converting between energy, nutrients and mass.     

Comparing pristine and depleted ecosystems: The Sørfjord, Norway versus the Gulf of St. Lawrence, Canada. Effects of intense fisheries on marine ecosystems

The Sørfjord, Norway, and the Gulf of St. Lawrence, Canada, are two sub-arctic ecosystems with similar trophic structure. However, in the Gulf of St. Lawrence, severe exploitation of groundfish stocks has lead to important shifts in the trophic structure. In the Sørfjord, the situation is different: fishing pressure is much lighter. Our hypothesis is that overexploitation leads to changes in the trophic structure and severely alters the resilience of ecosystems. Based on the same modelling approach (Ecopath with Ecosim) the food web structure was compared, using different ecosystem indicators. Patterns of food web structure and trophodynamics were contrasted. Cod was the keystone species in both ecosystems, and forage fish were also important. Even after similar environmental changes in both ecosystems, and after a reduction of fishing pressure in the Gulf of St. Lawrence, there is no recovery of cod stocks in this ecosystem. In the Sørfjord, after different perturbations (but not from the fishery), the ecosystem seems to return to its equilibrium.     

广西北部湾海域生态环境特征与保护对策

为促进广西北部湾海域生态环境的保护与可持续利用,文章根据2008—2018年广西北部湾海域生态环境和污染状况,分析广西北部湾海域生态环境变化趋势,并针对海洋生态环境存在的问题提出相应保护对策。研究结果表明:广西北部湾海域整体海水水质状况、典型海洋生态系统健康状况和海洋保护区生态环境状况均有所改善,沉积物质量保持良好且总体变化不大,入海污染物仍是主要污染源;结合广西海洋环境的特点,应加强陆海统筹和联防联控,加大生态修复资金的扶持力度,推进海洋生态修复,为广西开展海洋生态环境治理与修复提供技术支撑。     

Fatty acid transfer in the food web of a coastal Mediterranean lagoon: Evidence for high arachidonic acid retention in fish

The transfer of fatty acids (FAs) in the food web of a Mediterranean lagoon was studied using FA compositional patterns across several trophic levels. The structure of the food web was inferred from C and N stable isotopes values and an isotope mixing model was used in order to estimate the relative contribution of the different potential food sources to the biomass of consumers. Bidimensional plots of FA composition of food web components against their δ¹⁵N values indicated a general trend of increasing proportions of highly unsaturated fatty acids (HUFAs) with increasing trophic levels while the proportions of saturated fatty acids (SAFAs) and 18-carbon polyunsaturated fatty acids (PUFAs) decreased. Using the relative contributions of food sources to consumers and their FA compositions, a model was built in order to estimate the PUFA composition of consumer mixed diets which was compared to consumer PUFA profiles. The latter allowed the identification of the PUFAs which were mostly enriched/retained in consumer lipids. There was a surprisingly high retention of arachidonic acid (ARA), a trend which challenges the idea of low ARA needs in marine fish and suggests the important physiological role of this essential FA for fish in estuarine environments.     

Lower trophic levels and detrital biomass control the Bay of Biscay continental shelf food web: Implications for ecosystem management

The Bay of Biscay (North-East Atlantic) has long been subjected to intense direct and indirect human activities that lead to the excessive degradation and sometimes overexploitation of natural resources. Fisheries management is gradually moving away from single-species assessments to more holistic, multi-species approaches that better respond to the reality of ecosystem processes. Quantitative modelling methods such as Ecopath with Ecosim can be useful tools for planning, implementing and evaluating ecosystem-based fisheries management strategies. The aim of this study was therefore to model the energy fluxes within the food web of this highly pressured ecosystem and to extract practical information required in the diagnosis of ecosystem state/health. A well-described model comprising 30 living and two non-living compartments was successfully constructed with data of local origin, for the Bay of Biscay continental shelf. The same level of aggregation was applied to primary producers, mid-trophic-levels and top-predators boxes. The model was even more general as it encompassed the entire continuum of marine habitats, from benthic to pelagic domains. Output values for most ecosystem attributes indicated a relatively mature and stable ecosystem, with a large proportion of its energy flow originating from detritus. Ecological network analysis also provided evidence that bottom-up processes play a significant role in the population dynamics of upper-trophic-levels and in the global structuring of this marine ecosystem. Finally, a novel metric based on ecosystem production depicted an ecosystem not far from being overexploited. This finding being not entirely consistent over indicators, further analyses based on dynamic simulations are required.     

2019年中国近海海洋热浪特征研究

海洋热浪是指在一定海域内发生的海表温度异常偏高的现象,海洋热浪的发生会对海洋生态系统的功能与服务产生重大影响。近年来,中国近海海洋热浪频发,引起越来越多的关注。本文利用逐日海表温度观测资料对2019年中国近海海洋热浪特征进行研究分析,结果显示:2019年,在中国近海海南岛周边海域、长江口附近海域、渤海海域和江苏外海海域海洋热浪高发,频次为7～12次;北部湾附近海域海洋热浪发生时间最长,超过150 d。在此基础上,以北部湾海域2?4月经历的一次持续时间长达数月的海洋热浪事件为例,分析了事件期间对应的气温场、气压场以及风场的异常,并进一步分析了北部湾不同季节海洋热浪事件与各气象背景场的对应关系。总体而言,2019年北部湾海域2?4月海洋热浪期间,气温偏高,副热带高压面积偏大、位置偏西偏北,冬季风偏弱,是此海洋热浪发生与维持的重要原因。     

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.     

Food web analysis in two permanently open temperate estuaries: consequences of saltmarsh loss?

Saltmarsh vegetation, seston and microphytobenthos are all conspicuous components of most temperate estuaries and they potentially contribute to the estuarine food chain. Yet their relative contributions are unclear, as is the significance of saltmarsh losses through natural and human-induced impacts. This study aimed to quantitatively determine the contribution of various types of primary producers to detritus in the Walpole-Nornalup Estuary and Leschenault Inlet, two permanently open estuaries in SW Australia, and, estimate the flow of different types of detritus to higher trophic levels, using carbon ((13)C) and nitrogen ((15)N) stable isotopes as tracers. Results of the mixing model indicated that seston, microphytobenthos and to some extent seagrass and fringing saltmarsh were the main contributors to the detrital pool in both estuaries. However, the relative contribution of different primary producers varied both within and between estuaries. The contribution of saltmarsh was higher at sites close to rivers and dense fringing vegetation, while seston, microphytobenthos and seagrass dominated the detrital material at other sites. Benthic harpacticoid copepods were shown to feed on detritus though they appeared to actively select for different components of the detritus depending on site and estuary. Isotopic signatures of other consumers indicated that fish and invertebrates derived nutrients from MPB and detritus, either directly as food or indirectly through feeding on invertebrates. The overall contribution of saltmarsh to detritus was lower in Leschenault Inlet than in Walpole-Nornalup Estuary, possibly as a result of increased clearing of fringing vegetation around Leschenault Inlet. This pattern was however not reflected in harpacticoid food. Therefore, although losses of fringing saltmarsh around estuaries have the potential to significantly affect estuarine food webs, the significance of such losses will be site- and estuary-dependent.     

Carbon flows in the benthic food web at the deep-sea observatory HAUSGARTEN (Fram Strait)

The HAUSGARTEN observatory is located in the eastern Fram Strait (Arctic Ocean) and used as long-term monitoring site to follow changes in the Arctic benthic ecosystem. Linear inverse modelling was applied to decipher carbon flows among the compartments of the benthic food web at the central HAUSGARTEN station (2500 m) based on an empirical data set consisting of data on biomass, prokaryote production, total carbon deposition and community respiration. The model resolved 99 carbon flows among 4 abiotic and 10 biotic compartments, ranging from prokaryotes up to megafauna. Total carbon input was 3.78±0.31 mmol C m⁻² d⁻¹, which is a comparatively small fraction of total primary production in the area. The community respiration of 3.26±0.20 mmol C m⁻² d⁻¹ is dominated by prokaryotes (93%) and has lower contributions from surface-deposit feeding macro- (1.7%) and suspension feeding megafauna (1.9%), whereas contributions from nematode and other macro- and megabenthic compartments were limited to <1%. The high prokaryotic contribution to carbon processing suggests that functioning of the benthic food web at the central HAUSGARTEN station is comparable to abyssal plain sediments that are characterised by strong energy limitation. Faunal diet compositions suggest that labile detritus is important for deposit-feeding nematodes (24% of their diet) and surface-deposit feeding macrofauna (∼44%), but that semi-labile detritus is more important in the diets of deposit-feeding macro- and megafauna. Dependency indices on these food sources were also calculated as these integrate direct (i.e. direct grazing and predator–prey interactions) and indirect (i.e. longer loops in the food web) pathways in the food web. Projected sea-ice retreats for the Arctic Ocean typically anticipate a decrease in the labile detritus flux to the already food-limited benthic food web. The dependency indices indicate that faunal compartments depend similarly on labile and semi-labile detritus, which suggests that the benthic biota may be more sensitive to changes in labile detritus inputs than when assessed from diet composition alone. Species-specific responses to different types of labile detritus inputs, e.g. pelagic algae versus sympagic algae, however, are presently unknown and are needed to assess the vulnerability of individual components of the benthic food web.