Decadal changes in a NW Mediterranean Sea food web in relation to fishing exploitation

We analysed changes in the ecological roles of species, trophic structure and ecosystem functioning using four standardized mass-balance models of the South Catalan Sea (North-western Mediterranean). Models represented the ecosystem during the late 1970s, mid 1990s, early 2000s, and a simulated no-fishing scenario. The underlying hypothesis was that ecosystem models should quantitatively capture the increasing exploitation in the ecosystem from the 1970s to 2000s, as well as differences between the exploited and non-exploited scenarios. Biomass showed a general decrease, while there was an increase in biomass at lower trophic levels (TL) from the 1970s to 2000s. The efficiency of energy transfer (TE) from lower to higher TLs significantly increased with time. The ecosystem during the 1990s showed higher biomass and flows than during the 1970s and 2000s due to an increase in small pelagic fish biomass (especially sardines). Exploited food webs also showed similarities in terms of general structure and functioning due to high intensity of fishing already in the 1970s. This intensity was highlighted with low trophic levels in the catch, high consumption of production by fisheries, medium to high primary production required to sustain the catches and high losses in secondary production due to fishing. Significant differences on ecosystem structure and functioning were highlighted between the exploited and no-fishing scenarios. Biomass of higher TLs increased under the no-fishing scenario and the mean trophic level of the community and the fish/invertebrate biomass ratios were substantially lower in exploited food webs. The efficiency of energy transfer (TE) from lower to higher TLs was lower under the no-fishing scenario, and it showed a continuous decrease with increasing TL. Marine mammals, large hake, anglerfish and large pelagic fish were identified as keystone species of the ecosystem when there was no fishing, while their ecological importance notably decreased under the exploited periods. On the contrary, the importance of small-sized organisms such as benthic invertebrates and small pelagic fish was higher in exploited food webs.     

Trophic relationships in the recently impounded Bagré reservoir in Burkina Faso

The trophic dynamics of Bagré reservoir which has been recently impounded in Burkina Faso was based on the data collected during 1997–1998 period using the Ecopath model and software. Total fish biomass is 22.63 t km⁻² and mainly represents trophic levels (TLs) 2 and 3. The trophic food chain is relatively long and the overall transfer efficiency is quite low. Grazing foodweb based on primary producers is prominent in the reservoir ecosystem and detritus plays a less significant role. Seasonal and long-term variations in water quality have significant influences on the lower TLs clearly showing a bottom-up functioning of the ecosystem. Environmental degradations, such as siltation occurring in the lake, suggest possible risks in limiting ecosystem productivity.     

EcoTroph: Modelling marine ecosystem functioning and impact of fishing

EcoTroph (ET) is a model articulated around the idea that the functioning of aquatic ecosystems may be viewed as a biomass flow moving from lower to higher trophic levels, due to predation and ontogenetic processes. Thus, we show that the ecosystem biomass present at a given trophic level may be estimated from two simple equations, one describing biomass flow, the other their kinetics (which quantifies the velocity of biomass transfers towards top predators). The flow kinetic of prey partly depends on the abundance of their predators, and a top-down equation expressing this is included in the model. Based on these relationships, we simulated the impact on a virtual ecosystem of various exploitation patterns. Specifically, we show that the EcoTroph approach is able to mimic the effects of increased fishing effort on ecosystem biomass expected from theory. Particularly, the model exhibits complex patterns observed in field data, notably cascading effects and ‘fishing down the food web’. EcoTroph also provides diagnostic tools for examining the relationships between catch and fishing effort at the ecosystem scale and the effects of strong top-down controls and fast-flow kinetics on ecosystems resilience. Finally, a dynamic version of the model is derived from the steady-state version, thus allowing simulations of time series of ecosystem biomass and catches. Using this dynamic model, we explore the propagation of environmental variability in the food web, and illustrated how exploitation can induce a decrease of ecosystem stability. The potential for applying EcoTroph to specific ecosystems, based on field data, and similarities between EcoTroph and Ecopath with Ecosim (EwE) are finally discussed.     

Reconstructing historical marine ecosystems using food web models: Northern British Columbia from Pre-European contact to present

Mass-balance trophic models (Ecopath with Ecosim) are developed for the marine ecosystem of northern British Columbia (BC) for the historical periods 1750, 1900, 1950 and 2000 AD. Time series data are compiled for catch, fishing mortality and biomass using fisheries statistics and literature values. Using the assembled dataset, dynamics of the 1950-based simulations are fitted to agree with observations over 50 years to 2000 through the manipulation of trophic flow parameters and the addition of climate factors: a primary production anomaly and herring recruitment anomaly. The predicted climate anomalies reflect documented environmental series, most strongly sea surface temperature and the Pacific Decadal Oscillation index. The best-fit predator–prey interaction parameters indicate mixed trophic control of the ecosystem. Trophic flow parameters from the fitted 1950 model are transferred to the other historical periods assuming stationarity in density-dependent foraging tactics. The 1900 model exhibited an improved fit to data using this approach, which suggests that the pattern of trophic control may have remained constant over much of the last century. The 1950 model is driven forward 50 years using climate and historical fishing drivers. The resulting ecosystem is compared to the 2000 model, and the dynamics of these models are compared in a predictive forecast to 2050. The models suggest similar restoration trajectories after a hypothetical release from fishing.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Identifying 120 years of decline in ecosystem structure and maturity of Great South Bay, New York using the Ecopath modelling approach

Mass balanced models yield valuable information regarding ecological function and delivery of ecosystem services, but often rely on data collected well before many species were reduced to fractions of their original abundance. Lagoonal systems, such as Great South Bay (GSB), NY, sit on the interface of terrestrial and marine ecosystems and are prone to anthropogenic stressors but proximity to land also makes the presence of data regarding historic populations and structure more likely. To quantify over a century of ecosystem change, Ecopath models were developed for GSB at each of four time periods where commercial and scientific data exist: 1880s, 1930s, 1980s and 2000s. The results indicated that the GSB has experienced a decline in ecosystem maturity, loss of top keystone predators, a decline in connectivity to the ocean though the reduction of migratory species and increasing dominance of low trophic level organisms. These changes undermine the delivery of ecosystem services, increase conflicts over limited resources and suggest that present day restoration targets fail to recognize appropriate baselines. We discuss the role of stochastic events, which result in state changes that could be defined as regime shifts, and ecosystem connectivity to the long-term stability of lagoonal systems.     

Importance of jumbo squid Dosidicus gigas (Orbigny, 1835) in the pelagic ecosystem of the central Gulf of California

The Humboldt squid is an important predator in the pelagic ecosystem of the central Gulf of California and the commercial catch of this species has increased over the past decade, probable due to a decrease of several top predators (sharks, large pelagic fish and the marine mammals) and the optimal feeding conditions in this area. Its high abundance and important position in the pelagic food web was quantified through two trophic models of the pelagic ecosystem of the central Gulf of California. Models represented conditions in 1980 and 2002, to document the decadal changes in ecosystem structure and function. The models were composed of 18 functional groups, including marine mammals, birds, fish, mollusks, crustaceans, and primary producers. Model results show direct negative effects on principal prey groups such as myctophids and pelagic red crab and positive effects on sharks, marine mammals and specifically sperm whales. It thus appears that the jumbo squid has an important role in the ecosystem and plays a central part in the overall energy flow as main food item for most top predators, and due to its predation of organisms on lower tropic levels.     

Carbon-based balanced trophic structure and flows in the offshore Lake Ontario food web before (1987-1991) and after (2001-2005) invasion-induced ecosystem change

Replicate mass-balanced solutions to Ecopath models describing carbon-based trophic structures and flows were developed for the Lake Ontario offshore food web before and after invasion-induced disruption. The food webs link two pathways of energy and matter flow: the grazing chain (phytoplankton-zooplankton-fish) and the microbial loop (bacteria-protozoans) and include 19 species-groups and three detrital groups. Mass-balance was achieved by using constrained optimization techniques to randomly vary initial estimates of biomass and diet composition. After the invasion, production declined for all trophic levels and species-groups except Chinook salmon. The trophic level (TL) increased for smelt, adult sculpin, adult alewife and Chinook salmon. Changes to ecotrophic efficiencies indicate a reduction in phytoplankton grazing, increased predation pressure on Mysis, adult smelt and alewife and decreased predation pressure on protozoans. Specific resource to consumer TTE changed; increasing for protozoans (8.0-11.5%), Mysis (0.6-1.0%), and Chinook salmon (1.0-2.3%) and other salmonines (0.4-0.5%) and decreasing for zooplankton (20.2-15.1%), prey-fish (9.7-8.8%), and benthos (1.7-0.6%). Direct trophic influences of recent invasive species were low. The synchrony of the decline in PP and species-group production indicates strong bottom-up influence. Mass balance required an increase of two to threefold in lower trophic level biomass and production, confirming a previously observed paradoxical deficit in lower trophic level production. Analysis of food web changes suggest hypotheses that may apply to other similar large pelagic systems including, (1) as pelagic primary productivity declines, overgrazing of zooplankton results in an increase in protozoan production and a loss of trophic transfer efficiency, (2) habitat and food web changes increased Mysis predation on Diporeia and contributed to their recent decline, and (3) production of Chinook salmon, the primary piscivore, was uncoupled from pelagic production processes. This study demonstrates the value of food web models to better understand the impact of invasive species and to develop novel hypotheses concerning trophic influences.     

Ecological role of stomatopods (mantis shrimps) and potential impacts of trawling in a marine ecosystem of the southeast coast of India

Ecosystems are balanced by nature and each component in the system has a role in the sustenance of other components. A change in one component would invariably have an effect on others. Stomatopods (mantis shrimps) are common and ecologically important predatory crustaceans in tropical marine waters. The ecological role of mantis shrimps and potential impacts of trawling in a marine ecosystem were estimated using Ecopath with Ecosim (EwE) Version 5.0 software, by constructing a mass balanced Ecopath model of Parangipettai (Porto Novo) ecosystem. Based on fisheries information from the region, 17 ecological groups were defined including stomatopods. Both primary and secondary data on biomass, P/B, Q/B and diet composition were used as basic inputs. The mass balanced model gave a total system throughput of 14,756 t km⁻² year⁻¹. The gross efficiency of 0.000942 indicated higher contribution of lower food chain groups in the fishery though the mean trophic level was 3.08. The immature and developing stage of the ecosystem was indicated by the ratio of total primary production and total respiration (1.832) and the net system production (2643.30 t km⁻² year⁻¹). Key indices (flow to detritus, net efficiency and omnivory index), split mortality rates and mixed trophic impact of different ecological groups were obtained from the model. A flow diagram was constructed to illustrate the trophic interactions, which explained the biomass flows in the ecosystem with reference to stomatopods. Two temporal simulations were made, with 10 year durations in the mass balanced Ecopath model by using ecosim routine incorporated in EwE software. The effect of decrease in biomass of stomatopods in the ecosystem was well defined, in the first run with increase in stomatopod fishing mortality, and the group showed a high positive impact on benthopelagic fish biomass increase (129%). The simulation with increase in trawling efforts resulted in the biomass decline of different ecological groups as elasmobranchs to 1%, stomatopods to 2%, crabs and lobsters to 36%, cephalopods to 63%, mackerel to 78%, and shrimps to 89%. Present study warns stomatopod discards and further increase in trawling efforts in the region and it explained the need for ecosystem based fisheries management practices for the sustainability of marine fisheries.     

Exploring fisheries strategies for the western English Channel using an ecosystem model

An ecosystem model of the western English Channel ecosystem in 1994 was used to explore the effects of the use of a fishing policy optimization routine on profits, number of jobs and ecosystem structure. The optimization for single objective led to the specialization of the fishing fleet, with some fleet types being almost excluded. The profits and mainly the job optimizations led to big changes in the ecosystem structure, with loss of diversity, but the overall biomass of all vertebrate groups represented in the model increased considerably. For the objective focusing on ecosystem structure, there was an increase in biodiversity, with many long-lived groups predicted to increase, although the overall vertebrate biomass suffered just a small increase. An “ideal” mixed policy configuration was found when slightly greater weight was given to ecosystem structure than was given to profits and jobs. This scenario led to an overall reduction in effort but also to increased profits and biodiversity, while keeping the number of jobs at the same level as the baseline estimates. The results of the optimizations showed that the average trophic level of the catches is quite resistant to changes in the underlying system structure. On the other hand, despite the high level of aggregation of the model structure, a biodiversity index estimated by the model presented large changes as a function of the weights placed on the single policy functions, reflecting the changes in the system structure. The output of the application of the fishing optimization presented here should be considered in qualitative rather than in quantitative terms as an aid and part contribution to the complicated discussions on future long term management actions. Nonetheless it points to an overall reduction in fishing capacity, an objective widely accepted within the scientific community, while keeping the fishery in a profitable state.     

Assessment of impacts of invasive fishes on the food web structure and ecosystem properties of a tropical reservoir in India

A network model of trophic interactions in a tropical reservoir in India was developed with the objective to quantify matter and energy flows between system components and to study the impact of invasive fishes on the ecosystem. Structure of flows and their distribution within and between trophic levels were analysed by aggregating single flows into combined flows for discrete trophic levels. The trophic flows primarily occurred in the first four trophic level (TL) and the food web structure in this reservoir ecosystem was characterized by the dominance of low TL organisms, with the highest TL of only 3.57 for the top predator. Highest system omnivory index (SOI) was observed for indigenous catfishes (0.422), followed by the exotic fish Mozambique Tilapia (0.402). Nile Tilapia and Pearl spots show the highest niche overlap which suggests high competition for similar resources. The mixed trophic impact routine reveals that an increase in the abundance of the African catfish would negatively impact almost all fish groups such as Indian major carps, Pearl spots, indigenous catfishes and Tilapines. The other invasive fish Mozambique Tilapia adversely affects the indigenous catfishes. The most interesting observation in this study is that the most dominant invasive fish in this reservoir, the Nile Tilapia does not negatively impact any of the fish groups. In fact it positively impacts the Indian major carps. The direct and indirect effects of predation between system components (i.e. fish, invertebrates, phytoplankton and detritus) are quantitatively described and the possible influence and role in the ecosystem's functioning of the invasive fish species are discussed.     

Negative selection effects suppress relationships between bacterial diversity and ecosystem functioning

I assembled bacterial communities to explore the effects of bacterial diversity on multiple ecosystem functions, including bacterial community biovolume, decomposition of particulate organic matter, and biomass transfer to the next trophic level. The experiment used a two-way factorial design with four levels of bacterial diversity (one to four species) and the absence/presence of a bacterivorous ciliated protist Tetrahymena pyriformis as two main factors, and all possible combinations of the four bacterial taxa nested within each diversity level. Bacterial community biovolume increased as bacterial diversity increased, a result due largely to positive selection effects. Decomposition and consumer abundance, however, were unaffected by bacterial diversity, though both varied among bacterial composition treatments. Negative selection effects, the dominance of species that do not contribute significantly to ecosystem functioning, accounted for the lack of diversity effects on decomposition and consumer abundance. The presence of Tetrahymena reduced bacterial community biovolume but increased decomposition, without altering the diversity-functioning relationships. Decomposition was strongly linked with consumer abundance such that communities supporting larger consumer biomass exhibited higher decomposition rates. This study suggests that if the negative selection effect is common, as it might be when examining ecosystem variables other than biomass (due to the presence of keystone species that can contribute disproportionably to ecosystem functioning relative to their abundances), basic bacteria-mediated ecosystem processes, such as decomposition and energy transfer to the next trophic level, may not increase with bacterial diversity.     

Integrative trophic network assessments of a lentic ecosystem by key ecological approaches of water chemistry, trophic guilds, and ecosystem health assessments along with an ECOPATH model

The objective of this study was to describe the trophic structure and energy flow in a lentic ecosystem in South Korea. Physicochemical water conditions were evaluated along with the reservoir ecosystem health using a multimetric IBI model. Nutrient analyses of the reservoir showed a nutrient rich and hypereutrophic system. Guild analysis revealed that tolerant and omnivorous species dominated the ecosystem. Tolerant fish, as a proportion of the number of individuals, were associated (R² > 0.90, p < 0.01) with TN and TP, the key indicators of trophic state in lentic ecosystems. The mean Reservoir Ecosystem Health Assessment (REHA) score was 19.3 during the study, which was judged as in ‘fair to poor’ condition. A trophic analysis of the reservoir estimated by the ECOPATH model shows that most activity in terms of energy flow occurred in the lower part of the trophic web, where there was intensive use of primary producers as a food source. Consequently, of the 10 consumer groups, nine fell within trophic levels <2.8. Trophic levels (TL) estimated from the weighted average of prey trophic levels varied from 1.0 for phytoplankton, macrophytes, and detritus to 3.25 for the top predator, Pseudobagrus fulvidraco. Our integrated approach to trophic network analysis may provide a key tool for determining the effects of nutrient influx on energy flow pathways in lentic ecosystems.     

Assessing the carrying capacity of tilapia in an intertidal mangrove-based polyculture system of Pearl River Delta, China

A trophic model of an intertidal mangrove-based polyculture system in Pearl River Delta, China, was constructed using the Ecopath with Ecosim software. This polyculture system was chosen since it is the first integrated multi-trophic aquaculture (IMTA) system that was constructed on the basis of mangrove planting in China. The energy flows, ecosystem property, and carrying capacity of tilapia in the polyculture system were analyzed and evaluated. The results show the trophic level of 1.00 for primary producers and detritus to 2.85 for grass carp. The geometric mean of the trophic transfer efficiencies was 7.0%, with 7.2% from detritus and 6.8% from primary producers within the system. The ecosystem property indices show that this polyculture system has a high value of total primary production/total respiration (TPP/TR) and total primary production/total biomass (TPP/TB), together with low Finn's cycled index (FCI), Finn's mean path length (FML), and connectance index (CI), indicating that this system is at a development stage according to Odum's theory. The principal fish cultured in the system is tilapia, and mixed trophic impacts (MTI) show that tilapia has a marked impact on most compartments in this system, and the carrying capacity was found to be a tilapia culture biomass of 5.8 t ha⁻¹ in the system.     

Response of balanced network models to large-scale perturbation: Implications for evaluating the role of small pelagics in the Gulf of Maine

Exploring the response of an ecosystem, and subsequent tradeoffs among its biological community, to human perturbations remains a key challenge for the implementation of an ecosystem approaches to fisheries (EAF). To address this and related issues, we developed two network (or energy budget) models, Ecopath and Econetwrk, for the Gulf of Maine ecosystem. These models included 31 network “nodes” or biomass state variables across a broad range of trophic levels, with the present emphasis to particularly elucidate the role of small pelagics. After initial network balancing, various perturbation scenarios were evaluated to explore how potential changes to different fish, fisheries and lower trophic levels can affect model outputs. Categorically across all scenarios and interpretations thereof, there was minimal change at the second trophic levels and most of the “rebalancing” after a perturbation occurred via alteration of the diet matrix. Yet the model results from perturbations to a balanced energy budget fall into one of three categories. First, some model results were intuitive and in obvious agreement with established ecological and fishing theory. Second, some model results were counter-intuitive upon initial observation, seemingly contradictory to known ecological and fishing theory; but upon further examination the results were explainable given the constraints of an equilibrium energy budget. Finally, some results were counter-intuitive and difficult to reconcile with theory or further examination of equilibrium constraints. A detailed accounting of biomass flows for example scenarios explores some of the non-intuitive results more rigorously. Collectively these results imply a need to carefully track biomass flows and results of any given perturbation and to critically evaluate the conditions under which a new equilibrium is obtained for these types of models, which has implications for dynamic simulations based off of them. Given these caveats, the role of small pelagics as a prominent component of this ecosystem remains a robust conclusion. We discuss how one might use this approach in the context of further developing an EAF, recognizing that a more holistic, integrated perspective will be required as we continue to evaluate tradeoffs among marine biological communities.     

Effects of near‐future ocean acidification,fishing, and marine protection on a temperate coastal ecosystem

Understanding ecosystem responses to global and local anthropogenic impacts is paramount to predicting future ecosystem states. We used an ecosystem modeling approach to investigate the independent and cumulative effects of fishing, marine protection, and ocean acidification on a coastal ecosystem. To quantify the effects of ocean acidification at the ecosystem level, we used information from the peer‐reviewed literature on the effects of ocean acidification. Using an Ecopath with Ecosim ecosystem model for the Wellington south coast, including the Taputeranga Marine Reserve (MR), New Zealand, we predicted ecosystem responses under 4 scenarios: ocean acidification + fishing; ocean acidification + MR (no fishing); no ocean acidification + fishing; no ocean acidification + MR for the year 2050. Fishing had a larger effect on trophic group biomasses and trophic structure than ocean acidification, whereas the effects of ocean acidification were only large in the absence of fishing. Mortality by fishing had large, negative effects on trophic group biomasses. These effects were similar regardless of the presence of ocean acidification. Ocean acidification was predicted to indirectly benefit certain species in the MR scenario. This was because lobster (Jasus edwardsii) only recovered to 58% of the MR biomass in the ocean acidification + MR scenario, a situation that benefited the trophic groups lobsters prey on. Most trophic groups responded antagonistically to the interactive effects of ocean acidification and marine protection (46%; reduced response); however, many groups responded synergistically (33%; amplified response). Conservation and fisheries management strategies need to account for the reduced recovery potential of some exploited species under ocean acidification, nonadditive interactions of multiple factors, and indirect responses of species to ocean acidification caused by declines in calcareous predators.     

Representing mediating effects and species reintroductions in Ecopath with Ecosim

Ecosystem models play an important role in supporting ecosystem approaches to management. To improve the representation of how ecosystems work, ecosystem models should be able to represent mediating effects (e.g., habitat provision) that species provide to each other as well as species (re)introductions, both common situations that can strongly influence ecosystem dynamics. We examine how such processes can be incorporated into Ecopath with Ecosim (EwE), a widely used tool for represent aquatic ecosystems with the potential to support ecosystem-based management. We used the reintroduction of sea otters (Enhydralutris) to the west coast of Vancouver Island, British Columbia, Canada as a case study. The model demonstrates how to account for benefits provided by kelp forests by contributing to primary production, increased feeding areas and food availability through prey retention. It also demonstrates how the reintroduction and range expansion of sea otters can be represented in Ecospace, and the implications of these options.     

Modeling an exploited rocky coastal ecosystem: Bahia Tortugas, Mexico

A trophic structure model of the rocky coastal ecosystem in Bahia Tortugas, Mexico was constructed using Ecopath software to represent the main biomass flows in the system. Data for the model came from field observations (biomass estimates, stomach contents, and ecological observations for sea snails, abalones, lobster, some demersal finfishes, and macroalgae) carried out through ten field trips from 2006 to 2008. The results provide a snapshot of how the ecosystem operates. The model considers 23 functional groups. The total system throughput was 553 t/km²/year, 57% corresponds to internal consumption, 28% to respiration, 14% becomes detritus, and only 1% is removed through commercial fishing. The model suggests that even for exploited populations, predation and competition are heavier stresses than current fishing effort; however, because spiny lobster showed the second highest keystoneness’ index value, increasing fishing pressure on this group could strongly impact the entire ecosystem. We believe that this model has the potential to support management by allowing the exploration of the potential impacts of different fishing decisions at ecosystem level.