Which forcing factors fit? Using ecosystem models to investigate the relative influence of fishing and changes in primary productivity on the dynamics of marine ecosystems

Fishing mortality and primary production (or proxy for) were used to drive the dynamics of fish assemblages in 9 trophodynamic models of contrasting marine ecosystems. Historical trends in abundance were reconstructed by fitting model predictions to observations from stock assessments and fisheries independent survey data. The model fitting exercise derives values for otherwise unknown parameters that specify the relative strength of trophic interactions and, in some instances, a time series anomaly for changes in primary production. We measured how much better or worse were model predictions when bottom-up forcing by primary production were added to top-down forcing by fishing. Searching for cross system patterns, the relative contribution of fishing and changes in primary production, mediated through trophic interactions, are evaluated for the ecosystems as a whole and for selected similar species in different ecosystems. The analysis provides a simple qualitative way to explain which forcing factors have most influence on modeled dynamics. Both fishing and primary production forcing were required to obtain the best model fits to data. Fishing effects more strongly influenced 6 of 9 of the ecosystems, but primary production was more often found to be the main factor influencing the selected pelagic and demersal fish stock trends. Examination of sensitivity to ecological and model parameters suggests that the results are the product of complex food-web interactions rather than simple deterministic responses of the models.     

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.     

Two-way coupling versus one-way forcing of plankton and fish models to predict ecosystem changes in the Benguela

‘End-to-end’ models have been adopted in an attempt to capture more of the processes that influence the ecology of marine ecosystems and to make system wide predictions of the effects of fishing and climate change. Here, we develop an end-to-end model by coupling existing models that describe the dynamics of low (ROMS–N₂P₂Z₂D₂) and high trophic levels (OSMOSE). ROMS–N₂P₂Z₂D₂ is a biogeochemical model representing phytoplankton and zooplankton seasonal dynamics forced by hydrodynamics in the Benguela upwelling ecosystem. OSMOSE is an individual-based model representing the dynamics of several species of fish, linked through opportunistic and size-based trophic interactions. The models are coupled through a two-way size-based predation process. Plankton provides prey for fish, and the effects of predation by fish on the plankton are described by a plankton mortality term that is variable in space and time. Using the end-to-end model, we compare the effects of two-way coupling versus one-way forcing of the fish model with the plankton biomass field. The fish-induced mortality on plankton is temporally variable, in part explained by seasonal changes in fish biomass. Inclusion of two-way feedback affects the seasonal dynamics of plankton groups and usually reduces the amplitude of variation in abundance (top-down effect). Forcing and coupling lead to different predicted food web structures owing to changes in the dominant food chain which is supported by plankton (bottom-up effect). Our comparisons of one-way forcing and two-way coupling show how feedbacks may affect abundance, food web structure and food web function and emphasise the need to critically examine the consequences of different model architectures when seeking to predict the effects of fishing and climate change.     

Sensitivity of ecological models to their climate drivers: statistical ensembles for forcing.

Global and regional numerical models for terrestrial ecosystem dynamics require fine spatial resolution and temporally complete historical climate fields as input variables. However, because climate observations are unevenly spaced and have incomplete records, such fields need to be estimated. In addition, uncertainty in these fields associated with their estimation are rarely assessed. Ecological models are usually driven with a geostatistical model's mean estimate (kriging) of these fields without accounting for this uncertainty, much less evaluating such errors in terms of their propagation in ecological simulations. We introduce a Bayesian statistical framework to model climate observations to create spatially uniform and temporally complete fields, taking into account correlation in time and space, spatial heterogeneity, lack of normality, and uncertainty about all these factors. A key benefit of the Bayesian model is that it generates uncertainty measures for the generated fields. To demonstrate this method, we reconstruct historical monthly precipitation fields (a driver for ecological models) on a fine resolution grid for a climatically heterogeneous region in the western United States. The main goal of this work is to evaluate the sensitivity of ecological models to the uncertainty associated with prediction of their climate drivers. To assess their numerical sensitivity to predicted input variables, we generate a set of ecological model simulations run using an ensemble of different versions of the reconstructed fields. We construct such an ensemble by sampling from the posterior predictive distribution of the climate field. We demonstrate that the estimated prediction error of the climate field can be very high. We evaluate the importance of such errors in ecological model experiments using an ensemble of historical precipitation time series in simulations of grassland biogeochemical dynamics with an ecological numerical model, Century. We show how uncertainty in predicted precipitation fields is propagated into ecological model results and that this propagation had different modes. Depending on output variable, the response of model dynamics to uncertainty in inputs ranged from uncertainty in outputs that matched that of inputs to those that were muted or that were biased, as well as uncertainty that was persistent in time after input errors dropped.     

Investigations into a plankton population model: Mortality and its importance in climate change scenarios

The potential for marine plankton ecosystems to influence climate by the production of dimethylsulphide (DMS) has been an important topic of recent research into climate change. Several General Circulation Models, used to predict climate change, have or are being modified to include interactions of ecosystems with climate. Climate change necessitates that parameters within ecosystem models must change during long-term simulations, especially mortality parameters that increase as organisms are pushed toward the boundaries of their thermal tolerance. Changing mortality parameters can have profound influences on ecosystem model dynamics. There is therefore a pressing need to understand the influence of varying mortality parameters on the long-term behaviour of ecosystem models. This work examines the sensitivity of a model of DMS production by marine ecosystems to variations in three linear mortality coefficients. Significant differences in behaviour are observed, and we note the importance of these results in formulating ecosystem models for application in simulations of climate change.     

Long-term trends in the trophic structure of the North Sea fish community: evidence from stable-isotope analysis, size-spectra and community metrics

Fishing has wide-ranging impacts on marine ecosystems. One of the most pervasive signs of intensive fishing is "fishing down the food web", with landings increasingly dominated by smaller species from lower trophic levels. Decreases in the trophic level of landings are assumed to reflect those in fish communities, because size-selective mortality causes decreases in the relative abundance of larger species and in mean body size within species. However, existing analyses of fishing impacts on the trophic level of fish communities have focused on the role of changes in species composition rather than size composition. This will provide a biased assessment of the magnitude of fishing impacts, because fishes feed at different trophic levels as they grow. Here, we combine body size versus trophic level relationships for North Sea fishes (trophic level assessed using nitrogen stable-isotope analysis) with species-size-abundance data from two time-series of trawl-survey data (whole North Sea 1982-2000, central and northern North Sea 1925-1996) to predict long-term trends in the trophic structure of the North Sea fish community. Analyses of the 1982-2000 time-series showed that there was a slow but progressive decline in the trophic level of the demersal community, while there was no trend in the trophic level of the combined pelagic and demersal community. Analyses of the longer time-series suggested that there was no trend in the trophic level of the demersal community. We related temporal changes in trophic level to temporal changes in the slopes of normalised biomass size-spectra (which theoretically represent the trophic structure of the community), mean log₂ body mass and mean log₂ maximum body mass. While the size-based metrics of community structure showed long-term trends that were consistent with the effects of increased fishery exploitation, these trends were only correlated with trophic level for the demersal community. Our analysis suggests that the effects of fishing on the trophic structure of fish communities can be much more complex than previously assumed. This is a consequence of sampled communities not reflecting all the pathways of energy transfer in a marine ecosystem and of the absence of historical data on temporal and spatial changes in the trophic level of individuals. For the North Sea fish community, changes in size structure due to the differential effects of fishing on species and populations with different life histories are a stronger and more universal indicator of fishing effects than changes in mean trophic level.     

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.     

Mechanical clam dredging in Venice lagoon: ecosystem effects evaluated with a trophic mass-balance model

Harvesting of the invasive Manila clam, Tapes philippinarum, is the main exploitative activity in the Venice lagoon, but the mechanical dredges used in this free-access regime produce a considerable disturbance of the lagoon ecosystem. An ecosystem approach to study the complex effects of clam harvesting was implemented using a trophic mass-balance model. The trophic relations in the ecosystem were quantified with a mixed trophic impact analysis and further evaluated by considering different explanations for the " Tapes paradox", which consists of the apparent population enhancement of Manila clams by dredging and the apparent nutritional advantages that this species receives from re-suspended organic matter. The key-role played by this introduced species is highlighted by a network analysis that indicates a "wasp-waist control" of the system by Manila clams. The model constructed to characterise the present state of the Venice lagoon ecosystem is compared with models produced for a reconstructed past lagoon and a projected future lagoon. The future model was obtained by simulating the elimination of clam dredging in 10 years. The three different models were compared using thermodynamic and informational indices. Simulating the elimination of clam dredging produced a 33% increase in artisanal fishery catches, carried out by means of static gears, even with no change in fishing effort. These simulations also forecast an increase in the mean trophic level of the artisanal fishery catches as a positive effect of eliminating mechanical clam harvesting.     

The Scotia Sea krill fishery and its possible impacts on dependent predators: modeling localized depletion of prey

The nature and impact of fishing on predators that share a fished resource is an important consideration in ecosystem-based fisheries management. Krill (Euphausia superba) is a keystone species in the Antarctic, serving as a fundamental forage source for predators and simultaneously being subject to fishing. We developed a spatial multispecies operating model (SMOM) of krill-predator fishery dynamics to help advise on allocation of the total krill catch among 15 small-scale management units (SSMUs) in the Scotia Sea, with a goal to reduce the potential impact of fishing on krill predators. The operating model describes the underlying population dynamics and is used in simulations to compare different management options for adjusting fishing activities (e.g., a different spatial distribution of catches). The numerous uncertainties regarding the choice of parameter values pose a major impediment to constructing reliable ecosystem models. The pragmatic solution proposed here involves the use of operating models that are composed of alternative combinations of parameters that essentially try to bound the uncertainty in, for example, the choice of survival rate estimates as well as the functional relationships between predators and prey. Despite the large uncertainties, it is possible to discriminate the ecosystem impacts of different spatial fishing allocations. The spatial structure of the model is fundamental to addressing concerns of localized depletion of prey in the vicinity of land-based predator breeding colonies. Results of the model have been considered in recent management deliberations for spatial allocations of krill catches in the Scotia Sea and their associated impacts on dependent predator species.     

Key Features and Context‐Dependence of Fishery‐Induced Trophic Cascades

Abstract: Trophic cascades triggered by fishing have profound implications for marine ecosystems and the socioeconomic systems that depend on them. With the number of reported cases quickly growing, key features and commonalities have emerged. Fishery‐induced trophic cascades often display differential response times and nonlinear trajectories among trophic levels and can be accompanied by shifts in alternative states. Furthermore, their magnitude appears to be context dependent, varying as a function of species diversity, regional oceanography, local physical disturbance, habitat complexity, and the nature of the fishery itself. To conserve and manage exploited marine ecosystems, there is a pressing need for an improved understanding of the conditions that promote or inhibit the cascading consequences of fishing. Future research should investigate how the trophic effects of fishing interact with other human disturbances, identify strongly interacting species and ecosystem features that confer resilience to exploitation, determine ranges of predator depletion that elicit trophic cascades, pinpoint antecedents that signal ecosystem state shifts, and quantify variation in trophic rates across oceanographic conditions. This information will advance predictive models designed to forecast the trophic effects of fishing and will allow managers to better anticipate and avoid fishery‐induced trophic cascades.     

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.     

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.     

Linking species- and ecosystem-level impacts of climate change in lakes with a complex and a minimal model

To study the interaction between species- and ecosystem-level impacts of climate change, we focus on the question of how climate-induced shifts in key species affect the positive feedback loops that lock shallow lakes either in a transparent, macrophyte-dominated state or, alternatively, in a turbid, phytoplankton-dominated state. We hypothesize that climate warming will weaken the resilience of the macrophyte-dominated clear state. For the turbid state, we hypothesize that climate warming and climate-induced eutrophication will increase the dominance of cyanobacteria. Climate change will also affect shallow lakes through a changing hydrology and through climate change-induced eutrophication. We study these phenomena using two models, the full ecosystem model PCLake and a minimal dynamic model of lake phosphorus dynamics. Quantitative predictions with the complex model show that changes in nutrient loading, hydraulic loading and climate warming can all lead to shifts in ecosystem state. The minimal model helped in interpreting the non-linear behaviour of the complex model. The main output parameters of interest for water quality managers are the critical nutrient loading at which the system will switch from clear to turbid and the much lower critical nutrient loading – due to hysteresis – at which the system switches back. Another important output parameter is the chlorophyll-a level in the turbid state. For each of these three output parameters we performed a sensitivity analysis to further understand the dynamics of the complex model PCLake. This analysis showed that our model results are most sensitive to changes in temperature-dependence of cyanobacteria, planktivorous fish and zooplankton. We argue that by combining models at various levels of complexity and looking at multiple aspects of climate changes simultaneously we can develop an integrated view of the potential impact of climate change on freshwater ecosystems.     

Database-driven models of the world's Large Marine Ecosystems

We present a new methodology for database-driven ecosystem model generation and apply the methodology to the world's 66 currently defined Large Marine Ecosystems. The method relies on a large number of spatial and temporal databases, including FishBase, SeaLifeBase, as well as several other databases developed notably as part of the Sea Around Us project. The models are formulated using the freely available Ecopath with Ecosim (EwE) modeling approach and software. We tune the models by fitting to available time series data, but recognize that the models represent only a first-generation of database-driven ecosystem models. We use the models to obtain a first estimate of fish biomass in the world's LMEs. The biggest hurdles at present to further model development and validation are insufficient time series trend information, and data on spatial fishing effort.     

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.     

Human-driven spatial and temporal shift in trophodynamics in the Gulf of Maine,USA

Utilizing marine protected areas (MPAs) to isolate the ecological effects of human influence can help us understand our effect on systems and foster ecosystem-based approaches to management. Specifically, examining invertebrate prey community dynamics inside and outside an MPA may provide a measure of how altering human influence (i.e., certain fishing pressures) affects ecosystem interactions. We measured trophic interactions inside and outside a deep-water temperate MPA over 2 years. Predation rates on tethered, preferred groundfish prey (crabs) were initially identical inside and outside the MPA, but decreased outside the MPA after the commercial groundfish fishing season commenced. Predation trials using a ubiquitous prey species (brittle stars) and a less preferred prey species (urchins) served as controls, showing no MPA effect. Our experiments quantify differential predatory activity resulting from differences in human activity driven by an MPA, demonstrating important effects of fishing and regulations on the strength of trophic interactions.     

Parameter uncertainties in the modelling of vegetation dynamics—Effects on tree community structure and ecosystem functioning in European forest biomes

Dynamic vegetation models are useful tools for analysing terrestrial ecosystem processes and their interactions with climate through variations in carbon and water exchange. Long-term changes in structure and composition (vegetation dynamics) caused by altered competitive strength between plant functional types (PFTs) are attracting increasing attention as controls on ecosystem functioning and potential feedbacks to climate. Imperfect process knowledge and limited observational data restrict the possibility to parameterise these processes adequately and potentially contribute to uncertainty in model results. This study addresses uncertainty among parameters scaling vegetation dynamic processes in a process-based ecosystem model, LPJ-GUESS, designed for regional-scale studies, with the objective to assess the extent to which this uncertainty propagates to additional uncertainty in the tree community structure (in terms of the tree functional types present and their relative abundance) and thus to ecosystem functioning (carbon storage and fluxes). The results clearly indicate that the uncertainties in parameterisation can lead to a shift in competitive balance, most strikingly among deciduous tree PFTs, with dominance of either shade-tolerant or shade-intolerant PFTs being possible, depending on the choice of plausible parameter values. Despite this uncertainty, our results indicate that the resulting effect on ecosystem functioning is low. Since the vegetation dynamics in LPJ-GUESS are representative for the more complex Earth system models now being applied within ecosystem and climate research, we assume that our findings will be of general relevance. We suggest that, in terms of carbon storage and fluxes, the heavier parameterisation requirement of the processes involved does not widen the overall uncertainty in model predictions.     

Collapse and reorganization of a food web of Mwanza Gulf, Lake Victoria

Lake Victoria in East Africa is the world's second largest freshwater system. Over the past century the ecosystem has undergone drastic changes. Some 30 years after the introduction of Nile perch (Lates niloticus) and Nile tilapia (Oreochromis niloticus) in the 1950s, the highly diverse community of native haplochromines collapsed, leaving a system dominated by only four species: the native cyprinid dagaa (Rastrineobola argentea) and shrimp (Caridina nilotica), as well as the introduced Nile perch and Nile tilapia. More recently, an unexpected resurgence of haplochromines has been reported. To understand these changes in terms of ecosystem functioning and of changes in growth of trophic groups, we created mass balances of the food web near Mwanza, Tanzania, before, during, and after the Nile perch boom (1977, 1987, and 2005), using the application ECOPATH. We connected these mass balances with a dynamic model assuming linear trends in net growth rates of the trophic groups. Our analysis suggests that the Nile perch boom initially altered the biomass distribution over trophic levels. Also, results indicate that not only fishing but also changes at the detritivores' trophic level might have played an important role in driving changes in the system. Both the mass balances and the dynamic model connecting them reveal that, after a major distortion during the Nile perch boom, the biomass distribution over the main trophic levels had largely recovered its original (1977) state by 2005. However, no such return appeared in terms of community structure. Biodiversity in the new state is dramatically lower, consisting of introduced species and a few native surviving species. We conclude that at an aggregate level Lake Victoria's ecosystem has proved to be resilient in the sense that its overall trophic structure has apparently recovered after a major perturbation. By contrast, its intricate functional structure and associated biodiversity have proved to be fragile and seem unlikely to recover.     

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.     

A simulation model to explore the response of the Gulf of Maine food web to large-scale environmental and ecological changes

A dynamic simulation model was constructed using outputs from a balanced Gulf of Maine (GOM) energy budget model as the initial parameter set. The model was structured to provide a recipient control set of dynamics, largely based off of flows to and from different biological groups. The model was used to produce Monte Carlo simulations that were compared (percent change in biomass) with basecase simulations for a variety of scenarios. Changes in primary production, large increases in pelagic and demersal fish biomass, increases in fishing mortality, and large increases in top predators such as baleen whales and pinnepids were simulated. These scenarios roughly simulated the potential impacts of climate change, altered fishing pressure, additional protected species mitigations, and combinations thereof. Results suggest that the GOM system is primarily influenced by bottom-up processes involving phytoplankton, zooplankton, and bacterial biomass. Pelagic and demersal fish were important in determining trends in some of the scenarios. Marine mammals, large pelagic fish, and seabirds have a minor role in the GOM system in terms of biomass flows among the ecosystem components. The system is resilient to large-scale change due, in part to many predator–prey linkages. However, major alterations could occur from sustained climate change, high fishing rates, and by combinations of these types of external forcing mechanisms.