Impacts of Aggregation Methods and Trophospecies Number on the Structure and Function of Marine Food Webs

Aggregation of species with similar ecological properties is one of the effective methods to simplify food web researches.However,species aggregation will affect not only the complexity of modeling process but also the accuracy of models’outputs.Selection of aggregation methods and the number of trophospecies are the keys to study the simplification of food web.In this study,three aggregation methods,including taxonomic aggregation(TA),structural equivalence aggregation(SEA),and self-organizing maps(SOM),were analyzed and compared with the linear inverse model–Markov Chain Monte Carlo(LIM-MCMC)model.Impacts of aggregation methods and trophospecies number on food webs were evaluated based on the robustness and unitless of ecological net-work indices.Results showed that aggregation method of SEA performed better than the other two methods in estimating food web structure and function indices.The effects of aggregation methods were driven by the differences in species aggregation principles,which will alter food web structure and function through the redistribution of energy flow.According to the results of mean absolute percentage error(MAPE)which can be applied to evaluate the accuracy of the model,we found that MAPE in food web indices will increase with the reducing trophospecies number,and MAPE in food web function indices were smaller and more stable than those in food web structure indices.Therefore,trade-off between simplifying food webs and reflecting the status of ecosystem should be con-sidered in food web studies.These findings highlight the importance of aggregation methods and trophospecies number in the analy-sis of food web simplification.This study provided a framework to explore the extent to which food web models are affected by dif-ferent species aggregation,and will provide scientific basis for the construction of food webs.

An assessment of “fishing down marine food webs” in coastal states during 1950–2010

Mean trophic level of fishery landings（MTL） is one of the most widely used biodiversity indicators to assess the impacts of fishing. Based on the landing data compiled by Food and Agriculture Organization combined with trophic information of relevant species in Fish Base, we evaluated the status of marine fisheries from 1950 to 2010 for different coastal states in Pacific, Atlantic and Indian Oceans. We found that the phenomenon of ＂fishing down marine food webs＂ occurred in 43 states. Specifically, 27 states belonged to ＂fishing-through＂ pattern, and 16 states resulted from ＂fishing-down＂ scenario. The sign of recovery in MTL was common in the Pacific, Atlantic and Indian Oceans（occurred in 20 states）, but was generally accompanied by significantly decreased catches of traditional low trophic level species. In particular, 11 states showed significant declining catches of lower trophic levels. The MTL-based assessment of ＂fishing down marine food webs＂ needs to be interpreted cautiously.

Temporal variations of food web in a marine bay ecosystem based on LIM-MCMC model

Climate change has led to significant fluctuations in marine ecosystems,including alterations in the structure and function of food webs and ecosystem status.Coastal ecosystems are critical to the functioning of the earth’s lifesupporting systems.However,temporal variations in most of these ecosystems have remained unclear so far.In this study,we employed a linear inverse model with Markov Chain Monte Carlo(LIM-MCMC)combined with ecological network analysis to reveal the temporal variations of the food web in Haizhou Bay of China.Food webs were constructed based on diet composition data in this ecosystem during the year of 2011 and 2018.Results indicated that there were obvious temporal variations in the composition of food webs in autumn of 2011 and 2018.The number of prey and predators for most species in food web decreased in 2018 compared with 2011,especially for Trichiurus lepturus,zooplankton,Amblychaeturichthys hexanema,and Loligo sp.Ecological network analysis showed that the complexity of food web structure could be reflected by comprehensive analysis of compartmentalized indicators.Haizhou Bay ecosystem was more mature and stable in 2011,while the ecosystem’s self-sustainability and recovery from disturbances were accelerated from 2011 to 2018.These findings contribute to our understanding of the dynamics of marine ecosystems and highlight the importance of comprehensive analysis of marine food webs.This work provides a framework for assessing and comparing temporal variations in marine ecosystems,which provides essential information and scientific guidance for the Ecosystem-based Fisheries Management.

Impacts of species depletion on the food web structure of a marine ecosystem based on topological network analysis

Single-species management ignores the interactions between species,and ecosystem-based fisheries management(EBFM)has become a main method to fisheries management.Understanding food web structures and species interactions is essential for the implementation of EBFM and maintenance of ecosystem functions.Overfishing is one of the main reasons behind the depletion,which could even lead to the depletion of some target species in local areas.So understanding the impacts of species depletion on food web structures is important for the implementation of EBFM.The impacts of species depletion can be transmitted through the food web and cause the local extinction of both target and non-target species.In this study,topological network analysis was applied to examine the impacts of species depletion on the food web structure of Haizhou Bay.Results showed that fine crayfish Leptochela gracilis,squid Loligo sp.,and Japanese snapping shrimp Alpheus japonicus have the highest numbers of outgoing links(48,32 and 31 respectively);thus,these species may be considered key prey species.Whitespotted conger Conger myriaster,fat greenling Hexagrammos otakii,and bluefin gurnard Chelidonichthys kumu were key predators with the highest number of incoming links(37,36 and 35 respectively).The competition graphs derived from the Haizhou Bay food web were highly connected(more than 40%predators sharing over 10 common prey species),and showed close trophic interaction between high trophic level fishes.Simulation analysis showed that the food web structure has small changes to the depletion of species in a highly complex food web.The most-connected target species did not necessarily indicate high structural importance;however,some species with low connectivity may demonstrate stronger trophic interactions and play important ecological roles in the food web.But most species were more sensitive to the depletion of the most-connected target species than other target species(for instance,for zooplankton,closeness centrality 13.876 in D6,but closeness centrality 82.143 in original food web).Therefore,EBFM should focus on the most-connected target species,but also on those species with few but strong links and feeding relationships in the food web.

Determinant variables on the isotopic values of particulate organic matter in a neotropical floodplain

Particulate organic matter(POM)is an important energy source for aquatic consumers,understanding its origin and composition is essential for understanding the energetic dynamics of aquatic environments.The aim of this study was to analyze the relationship between POM and phytoplankton(isotopic values and chlorophyll concentration)and abiotic variables during dry and rainy seasons.Sampling was conducted in rivers and lagoons in the floodplain of the Upper ParanáRiver.We found a greater difference in ^(δ13)C values of POM between sampling points than between seasons,indicating that the composition of regional sources influences the composition of POM more than dry and rainy seasons.In addition,the concentration of chlorophyll during the dry season was positively correlated with ^(δ13)C values during that rainy period.Additionally,we found a relationship between factors limiting the growth of phytoplankton and ^(δ13)C values of POM,such as phosphate ions,indicating that variables that regulate phytoplankton growth tend to influence the composition of POM in river floodplains.Therefore,maintaining the variables that regulate the phytoplankton community is of fundamental importance for the composition of POM,an important energy source in aquatic environments.

Evaluation of the “fishing down marine food web” process in the north-west of Persian Gulf(Khuzestan Province) during the period of 2002–2011

The worldwide increase in commercial fisheries and its impact on ecosystems as well as inefficient fishery management have led to overfishing and frequent breakdown of traditional fish stocks. In this context, an analysis of Khuzestan inshore fisheries data covering the years 2002-2011, was conducted in reliance on testing for occurrence of the fishing down marine food webs （FDMFW） phenomenon in the North of Persian Gulf Large Marine Ecosystem （LME）. In this study, the mean trophic level （mTL） and the fishing-in-balance （FIB）-index of Khuzestan landings during this period of time were estimated using the trophic level of 47 fishery resources. Increase in total landings （I1） was observed, which explained the high fishing yield in major fishery resources （especially demersal）. Moreover, the moderates decreasing trend in mTL per decade, and the increasing trend in FIB-index were observed. The status of fishery resources in Khuzestan inshore waters （under exploited but not overexploited）, the rise in Y, FIB and slightly drop in mTL can be considered as indirect indicators of the fishing impacts on the trophic structure of marine communities. Based on this result, probability occurrence of FDMFW process in Khuzestan inshore waters is low to some extent. However, we suggest that the goal of management programs in Khuzestan inshore waters should prevent the continuance of this trend in the long-term using an ecosystem-based approach.

Topology-based analysis of pelagic food web structure in the central and eastern tropical Pacific Ocean based on longline observer data

The tropical Pacific Ocean supports many productive commercial fisheries.However,few studies of ecosystem structure in the tropical Pacific Ocean have been carried out.In this study,we analyzed the food web structure in the central and eastern tropical Pacific Ocean based on trophic relationships of 35 pelagic species collected by Chinese tuna longline observers from June to November in 2017.Topology indices(node degree,D;centrality indices,BC and CC;topological importance indices,TI^1,TI^3;keystone indices,K,K_t and K_b)and Key-Player algorithms(KPP-1,KPP-2)were used to select key species and construct a simplified food web combined with body size data.The Kendall rank correlation and hierarchical clustering analysis indicated that different topology indices resulted in consistent rankings of key species.Most key species were the same as those selected in other studies in the Pacific Ocean,such as Shortbill spearfish(Tetrapturus angustirostris),Swordfish(Xiphias gladius),Albacore tuna(Thunnus alalunga),cephalopods and scomber.The food web would be separated into many unconnected parts(F=0.632,FD=0.795,R^D=0.957)after the removal of the five key species,indicating the key roles of these species in the food web structure and stability.Body size was considered an influential indicator in constructing the simplified food web.This study can improve our understanding of the food web structure in the tropical Pacific Ocean and provide scientific basis for further ecosystem dynamics studies.

Spatial variations in food web structures with alternative stable states:evidence from stable isotope analysis in a large eutrophic lake

Food web structures are well known to vary widely among ecosystems. Moreover, many food web studies of lakes have generally attempted to characterize the overall food web structure and have largely ignored internal spatial and environmental variations. In this study, we hypothesize that there is a high degree of spatial heterogeneity within an ecosystem and such heterogeneity may lead to strong variations in environmental conditions and resource availability, in turn resulting in dif ferent trophic pathways. Stable carbon and nitrogen isotopes were employed for the whole food web to describe the structure of the food web in different sub-basins within Taihu Lake. This lake is a large eutrophic freshwater lake that has been intensively managed and highly influenced by human activities for more than 50 years. The results show significant isotopic dif ferences between basins with dif ferent environmental characteristics. Such differences likely result from isotopic baseline dif ferences combining with a shift in food web structure. Both are related to local spatial heterogeneity in nutrient loading in waters. Such variation should be explicitly considered in future food web studies and ecosystem-based management in this lake ecosystem.

A preliminary study on benthos food web structure oftidal zone in the Laoshan Bay by using stable carbon isotopes

By anayses of carbon isotopic composition (δ13C values) of the benthos collected in the Laoshan Bay in August 1993 and February and May 1994, it is found that the main food sources of the benthos can be divided into four groups in terms of carbon isotope composition: Particulate Organic Matter (POM), benthic diatoms, benthic macroalgae and the organic matter in sediments. The results show that the carbon isotopic composition of the benthic animals has a close relation with that of the food they take in. The Carbon isotopic compositions of benthos may be useful in elucidating their food sources. The carbon isotopic data have confirmed that POM is the main food source of the benthic filter feeders such as bivalves; crustaceans have a wide range of δ13C values, showing their food source has diversity; benthic diatoms are an important fraction of the food for most of gastropods. A preliminary investigation of the benthic-pelagic coupling in that region using the stable carban isotopic tracers has confirmed the importance of POM as a food source for benthos in this region, but the organic matter in sediments and benthic diatoms are also relatively important for a lot of benthic animals. The benthic-pelagic coupling in the Laoshan Bay temperate ecosystem is not so tight as that in ecosystems at higher latitudes such as northeastern water polynya.

Food web structure and trophic levels in a saltwater pond sea cucumber and prawn polyculture system

The food sources of aquacultured Apostichopus japonicus and the trophic levels of organisms in a sea cucumber（A. japonicus） and prawn（Penaeus japonica） polyculture system in a saltwater pond in Zhuanghe, Liaoning Province were examined using carbon and nitrogen stable isotopes. Across organisms, δ13C ranged from（–25.47±0.20）‰ to（–16.48±0.17）‰（mean±SD）, and δ15N ranged from（4.23±0.49）‰ to（12.44±0.09）‰. The δ13C and δ15N contents of A. japonicus, P. japonica and Fenneropenaeus chinensis were comparatively higher than those of other organisms. Values of δ13C and δ15N revealed that P. japonica, Hemigrapsus sanguineus and Neomysis japonica comprised the largest component of the diet of A. japonicus. The mean trophic level of the organisms in this saltwater pond polyculture system was（2.75±0.08）. P. japonica, A. japonicus, F. chinensis,Synechogobius hasta and Neomysis japonica were in the 3rd trophic level（2–3）; jellyfish, H. sanguineus and zooplankton were in the 2nd trophic level（1–2）; and Enteromorpha prolifera, benthic microalgae, periphyton and suspended matter primarily consisting of phytoplankton, bacteria and humus were in the primary trophic level（0–1）.

Stable isotopes in aquatic food web of an artificial lagoon in the Hangzhou Bay,China

Stable isotope values, δ13C and δ15N, were determined for four primary producers and 19 dominant consumers in a small artificial lagoon located in Hangzhou Bay. Based on these results the major pathways for energy flow and trophic structure of the artificial lagoon ecosystem were characterized. The mean δ13C values for the 19 consumers ranged from -22.99‰ to -14.24‰. Apart from so-iny mullet Liza haematocheila, the other 18 consumers had intermediate δ13C values between those of epibenthic microalgae and particulate organic matter (POM). The results of a multiple source linear mixing model (IsoSource model) indicated that 50% or more of the organic carbon in the tissues of most consumers was derived from epibenthic microalgae. This indicated that these primary producers were the main food source fueling the lagoon food web. The mean δ15N values for the 19 consumers varied between 4.93‰ and 12.97‰ and indicated four trophic levels in the lagoon. Four macroinvertebrates and zooplankton represented the primary consumers, whilst the other 14 consumers occupied the secondary and tertiary consumer levels. The 19 consumers were divided into three trophic guilds (detritivores/suspension feeders, omnivores and carnivores).

Construction of Food Web Model for Liusha Bay-Based on Stable Isotope Analysis

To investigate the food web structure of Liusha Bay,the δ13C and δ15N values of various organisms collected in Liusha Bay were measured via stable isotope technique.Based on the calculated δ15N value,the correlation between stable isotope ratio and trophic level were established to predict the trophic levels of shrimps,crabs,cephalopods and fishes.By comparing the analysis results of stomach contents of 24 organisms,two methods gave a similar consistence in approximately 71% of organisms within a 0.5 trophic level.It is concluded that stable isotope technique could be an effective method for studying ocean food web trophic level.

Removal of Agricultural Non-Point Source Pollutants by Artificial Aquatic Food Web System: A Study Case of the Control of Cynobacterial Bloom in Jiyu River

An artificial aquatic food web (AAFW) system was designed to remove the non-point source pollutants in eutrophic Jiyu river. A certain amount of Scenedesmus obliquus and Daphnia pulex was cultured in the system for the control of serious cyanobacterial bloom. The AAFW system was a continuous-flow system including one storage basin of 3 m3 capacity with polluted river water (the total nitrogen-TN: 4.49 mg⋅l-1;the total phosphorus-TP: 0.192 mg⋅l-1), one phytoplankton tank of 3 m3 capacity with an initial concentrations of S. obliquus about 5.8 × 103 ind⋅l-1, and one zooplankton growth chamber of 1.5 m3 capacity with an initial abundance of D. pulex about 22.5 ind⋅l-1. The system was optimized by setting hydraulic retention time of phytoplankton tank as 5 days and the experiments were operated for 45 days. Compared with the polluted river, TN and TP were removed about 28% and 47% by the AAFW system, respectively. The biomass of phytoplankton decrease from 6.33 mg⋅l-1 to 1.48 mg⋅l-1 and the percentage of cyanobacteria decrease from 43.93% to 2.36%, the biomass of Crustacean zooplankton increase from 0.34 mg⋅l-1 to 1.53 mg⋅l-1 and the percentage of D. pulex increase from 19.19% to 57.62%. Our results indicated that the AAFW system not only is an efficient, flexible system for reducing nutrient levels in tributary rivers, but also has an ability to control the cyanobacteria bloom and rebuilding the aquatic ecosystem from the polluted river water.

Leaf phenology rather than mycorrhizal type regulates soil nematode abundance,but collectively affects nematode diversity in seven common subtropical tree species

The underlying mechanisms of the relationships between tree species and the soil micro-food web in forest ecosystems remain uncertain,primarily ascribed to an insufficient understanding on how tree functional traits drive soil nematode communities,including in subtropical forests.We investigated the impacts of seven subtropical tree species(evergreen:Pinus massoniana,Mytilaria laosensis,Ilex chinensis,Michelia macclurei;and deciduous:Liquidambar formosana,Quercus acutissima,and Betula luminifera)on the soil nematode communities.We found that the abundance of soil nematodes was not affected by mycorrhizal types,but it was around 83%higher under the deciduous trees than the evergreen trees,indicating the importance of leaf phenology to the abundance of soil nematodes.Nonetheless,both the evergreen and the arbuscular mycorrhizal trees increased soil nematode diversity,resulting from changes in root traits and soil properties.Furthermore,root traits(root C,root N,and root C:N ratio),and soil properties(total C,total N,moisture content,and bulk density)were the best predictors of the community composition of soil nematodes,indicating a key role of resource quality and soil microhabitat in regulating soil nematodes.In contrast,the ectomycorrhizal trees had lower plant parasite and Wasilewska indices,and evenness,whereas the evergreen trees slightly improved the evenness of soil nematodes.This study suggests that tree species affect the soil food web through changes in soil conditions and plant functional traits in subtropical forests.

Effects of tree diversity on insect herbivory

Tree diversity has long been considered a key driver of insect herbivory in forest ecosystems.However,studies have given contradictory results:increased tree diversity can have positive,negative or neutral effects on insect herbivory.Since many issues can complicate the tree-herbivore interactions,the descriptor‘tree diversity’per se actually has only limited explanatory power for insect herbivory.Particularly,in addition to the direct bottom-up effects on insect herbivores,tree diversity may have stronger indirect top-down effects via natural enemies of insect herbivores.However,most research has addressed only direct bottom-up impacts.In fact,insect herbivory is the result of complex interactions(food webs)among all the species in a community.Although it is hard to integrate all functionally important species and dynamic information into food webs,the effects of tree diversity on insect herbivory are highly dependent on the dominant species at different trophic levels.It is important and practical to simultaneously consider the characteristics of main trees,herbivores,and natural enemies when attempting to predict the overall effects of forest tree diversity on insect herbivory.In order to ensure comparability between studies,it is necessary to classify them according to the descriptors of insect herbivory and tree diversity,and to compare results within each category.These measures might enhance our understanding of the mechanisms by which tree diversity drives insect herbivory and,in turn,help to develop sustainable pest management strategies for forests.

Construction and analysis of a coral reef trophic network for Qilianyu Islands,Xisha Islands

Qilianyu Islands coral reefs(QICR),located in the northeastern part of the South China Sea,has been affected by human activities and natural disturbance.To characterize the trophic structure,ecosystem properties and keystone species of this region,a food-web model for the QICR is developed using methods involving a mass-balance approach with Ecopath with Ecosim software.Trophic levels range from 1.00 for detritus and primary producers to 3.80 for chondrichthyes.The mean trophic transfer efficiency for the entire ecosystem is 13.15%,with 55%of total energy flow originating from primary producers.A mixed trophic impact analysis indicates that coral strongly impacts most components of this ecosystem.A comparison of our QICR model with that for other coral reef ecosystems suggests that the QICR ecosystem is immature and/or is degraded.

Trophic relationships in the Changjiang River estuarine salt marshes: preliminary investigation from δ^(13)C and δ^(15)N analysis

To obtain information on food web structure in salt marshes of the Changjiang （Yangtze） River Estuary, the （ δ3C and δ15N values of primary producers and consumers were determined. The mean δ13C values of 31 dominant consumers ranged from -23.13‰ to -14.37‰. Except for several species （ Eriocheir sinensis, Sinonovacula constricta and Potamocorbula ustulata）, consumers had interme- diate δ13C values between those of benthic microalgae and Spartina alterniflora. The mean δ15N values of 31 dominant consumers varied between 6.87‰ and 13.33‰, which indicate three trophic levels in salt marshes of the Changjiang River Estuary. A total of 18 macroinvertebrates species and four fish species represented primary consumers with trophic levels ranging from 2.0 to 2.7. Secondary consumers included two maeroinvertebrates and seven fishes with trophic levels varying between 3.0 and 3.9. The consumers were divided into three trophic guilds, i.e., detritivorous/algae feeders, omnivores and carnivores. The detrital food chain was the main energy flow pathway in the salt marsh food web of the Changjiang River Estuary, and the marsh vascular plants were at least as equally important as microphytobenthos for secondary production. The important trophic function of the salt marsh habitats in the estuary is revealed.

Modeling Study of Seasonal Variation of the Pelagic-Benthic Ecosystem Characteristics of the Bohai Sea

Based on experiment data of the Sino-German comprehensive investigations in the Bohai Sea in 1998 and 1999, a simple coupled pelagic-benthic ecosystem multi-box model is used to simulate the ecosystem seasonal variation. The pelagic sub-model consists of seven state variables： phytoplankton, zooplankton, TIN, TIP, DOC, POC and dissolved oxygen （DO）. The benthic sub-model includes macro-benthos, meiobenthos, bacteria, detritus, TIN and TIP in the sediment. Besides the effects of solar radiation, water temperature and the nutrient from sea bottom exudation, land-based inputs are considered. The impact of the advection terms between the boxes is also considered. Meanwhile, the effects of the micro- bial-loop are introduced with a simple parameterization. The seasonal variations and the horizontal distributions of the ecosystem state variables of the Bohai Sea are simulated. Compared with the observations, the results of the multi-box model are reasonable. The modeled results show that about 13% of the photosynthesis primary production goes to the main food loop, 20% transfers to the benthic domain, 44% is consumed by the respiration of phytoplankton, and the rest goes to DOC. Model results also show the importance of the microbial food loop in the ecosystem of the Bohai Sea, and its contribution to the annual zooplankton production can be 60%-64%.

Spatial and temporal characteristics of trophic structure of the southern Huanghai Sea fish communities

The spatial and temporal characteristics of trophic structure of fish communities in the southern Huanghai Sea were examined based on the data sampled from bottom trawl surveys conducted during the autumn of 2000 and the spring of 2001. Hierarchical agglomerative cluster method and bootstrap randomization were used to identify significant trophic groups for each fish assemblage in the southern Huanghai Sea. A total of six major trophic groups were identified within this system, which classified predators based upon location in the water column or prey size （ i. e. , benthic to pelagic predators or fish to small invertebrate prey predators）. The similarity level used to identify significant trophic groups in each assemblage ranged from 24% to 34%. Although planktivores were the dominant trophic group in each assemblage （60% - 79% ）, there were spatial and temporal variations in the trophic structure, which reflected the differences in the abundance and availability of dominant preys. Simplified food webs were constructed to evaluate the most important trophic relationships between the dominant prey taxa and the fishes in each assemblage within this system. Although there were some differences in the key prey species among different food webs, pelagic prey items （mainly euphausiids and copepods） represent the most important energetic link between primary producers and higher trophic level predators. The trophic level for most fishes was between 3 and d, and the weighted mean trophic level for each assemblage ranged from 3.3 to 3.4. Compared with previous study in the mid-1980s, there was an obvious downward trend in the trophic level for most fish species, which resulted mainly from the fluctuation in key prey species in the Huanghai Sea. The decrease in the importance of Japanese anchovy seems to be offset by other abundant prey species such as Euphausia pacifica and copepods （ mainly Calanus sinicus ） .

Size fraction of phytoplankton and the contribution of natural plankton to the carbon source of Zhikong scallop Chlamys farreri in mariculture ecosystem of the Sanggou Bay

The biomass and size fraction of phytoplankton in terms of chlorophyll a（Chl a） was measured during four cruises conducted in April, July, October 2013 and January 2014 in mariculture area, the Sanggou Bay, China.Results show that total Chl a levels in the surface seawater of the Sanggou Bay generally range from 0.10 to 20.46μg/L, with an average value of 2.13 μg/L. Nano-phytoplankton was the most important size-fraction and accounted for about 65.1% of total Chl a. In order to evaluate the importance of the ＂protozoan trophic link＂ for energy transfer from the microbial loop to filter-feeding feeders, Zhikong scallop Chlamys farreri was then offered a natural planktonic community as potential prey. Results show that scallops obtained carbon source from natural plankton with the rate of 11 033.05 μg/（g·d）. Protists（nanoflagellates and ciliates） were the dominant source of carbon retained by scallop（48.78%）. The microbial loop provided 58.45% of the carbon source for farmed scallops. These results indicate that the microbial loop represent a valuable trophic resource in mariculture system of the Sanggou Bay.