Lynne J. Shannon

Impacts of Fishing Low-Trophic Level Species on Marine Ecosystems

High harvest levels of low–trophic level fishes may have cascading marine ecosystem effects. Low–trophic level species account for more than 30% of global fisheries production and contribute substantially to global food security. We used a range of ecosystem models to explore the effects of fishing low–trophic level species on marine ecosystems, including marine mammals and seabirds, and on other commercially important species. In five well-studied ecosystems, we found that fishing these species at conventional maximum sustainable yield (MSY) levels can have large impacts on other parts of the ecosystem, particularly when they constitute a high proportion of the biomass in the ecosystem or are highly connected in the food web. Halving exploitation rates would result in much lower impacts on marine ecosystems while still achieving 80% of MSY.

Global Seabird Response to Forage Fish Depletion—One-Third for the Birds

One-third of maximum fish biomass must be available for seabirds to sustain high breeding success. Determining the form of key predator-prey relationships is critical for understanding marine ecosystem dynamics. Using a comprehensive global database, we quantified the effect of fluctuations in food abundance on seabird breeding success. We identified a threshold in prey (fish and krill, termed “forage fish”) abundance below which seabirds experience consistently reduced and more variable productivity. This response was common to all seven ecosystems and 14 bird species examined within the Atlantic, Pacific, and Southern Oceans. The threshold approximated one-third of the maximum prey biomass observed in long-term studies. This provides an indicator of the minimal forage fish biomass needed to sustain seabird productivity over the long term.

End-To-End Models for the Analysis of Marine Ecosystems: Challenges, Issues, and Next Steps

Abstract There is growing interest in models of marine ecosystems that deal with the effects of climate change through the higher trophic levels. Such end-to-end models combine physicochemical oceanographic descriptors and organisms ranging from microbes to higher-trophic-level (HTL) organisms, including humans, in a single modeling framework. The demand for such approaches arises from the need for quantitative tools for ecosystem-based management, particularly models that can deal with bottom-up and top-down controls that operate simultaneously and vary in time and space and that are capable of handling the multiple impacts expected under climate change. End-to-end models are now feasible because of improvements in the component submodels and the availability of sufficient computing power. We discuss nine issues related to the development of end-to-end models. These issues relate to formulation of the zooplankton submodel, melding of multiple temporal and spatial scales, acclimation and adaptation, behavioral movement, software and technology, model coupling, skill assessment, and interdisciplinary challenges. We urge restraint in using end-to-end models in a true forecasting mode until we know more about their performance. End-to-end models will challenge the available data and our ability to analyze and interpret complicated models that generate complex behavior. End-to-end modeling is in its early developmental stages and thus presents an opportunity to establish an open-access, community-based approach supported by a suite of true interdisciplinary efforts.

The functioning of marine ecosystems: a fisheries perspective.

There is considerable evidence that environmental variability plays a major role in controlling abundance and distribution of marine populations and that fisheries alter ecosystem functioning and state. This overviewdocuments emergent, i.e. visible to us as observers, ecosystem-level ecological patterns and addresses important questions regarding the exploitation of marine resources. Do marine ecosystems function differently from terrestrial systems? Do multiple stable states exist in marine ecosystems? Does removal of top predators in marine ecosystems result in fundamental changes in the plankton communities (top-down ‘trophic cascades’), as observed in lakes? Alternatively, are marine ecosystems characterized by bottom-up control such that fishing predatory fish does not disturb community structure and function? Does heavy exploitation of forage species, such as anchovies and sardines, cause changes in the functioning of upwelling ecosystems? The key to answering these questions and exploring whether general principles apply lies in understanding the energy flow within the ecosystems. The chapter reviews different types of energy flow in marine ecosystems, i.e. bottom-up control (control by primary producers), top-down control (control by predators) and wasp-waist control (control by numerically dominant species). No general theory can yet be ascribed to the functioning of marine ecosystems. Ecological understanding and models of ecosystem functioning are provisional and subject to change, and common sense is not sufficient when studying complex dynamic systems. However, tentative and partial generalizations are proposed, namely that bottom-up control predominates; top-down control plays a role in dampening ecosystem-level fluctuations; trophic cascades seldom occur; and wasp-waist control is most probable in upwelling systems. Moreover, alternation and large-scale synchronized fluctuations in fish stocks, stability of fish communities and emergent features such as size spectra are potentially important patterns when assessing states and changes in marine ecosystems. New and meaningful indicators, derived from our current understanding of marine ecosystem functioning, can be used to assess the impact of fisheries and to promote responsible fisheries in marine ecosystems.

Trophic flows in the southern Benguela during the 1980s and 1990s

Mass-balanced models of trophic flows in the southern Benguela ecosystem suggest a 10% increase in zooplankton biomass between the 1980s and the 1990s, in agreement with observed trends of increased zooplankton abundance off South Africa over the last few decades. Minimum hake biomass in balanced trophic models is substantially larger than survey and other model estimates,suggestingundersamplingofhakesinsurveysandunderestimationofjuvenilehakemortality.Modelbiomassandmean annual production of five important small pelagic fish groups were larger in the 1990s, and total catches were smaller than in the 1980s. Estimates of biomass per trophic level, transfer efficiencies, mixed trophic impacts and many other ecosystem attributes suggest that trophic functioning of the southern Benguela ecosystem was similar in the 1980s and 1990s. Because catches were lowerandmodelzooplanktonandsmallpelagicfishbiomasseswerelargerinthe1990s,theecosystemwaslesstightlyconstrained by predators (including fishers) and food availability than in the 1980s. Fishing took place at low trophic levels compared to other systems. Despite smaller total catches in the 1990s, fishing was ecologically more expensive (from higher trophic levels) during the 1990s than in the 1980s because snoek and hake catches were large. There was greater shared niche overlap of small pelagic fish predators in the 1990s than in the 1980s. Mean transfer efficiency was 12%. Transfer of biomass at trophic levels III–V appears to be more efficient in the southern Benguela than in other upwelling ecosystems. Primary production required to sustain catches in the southern Benguela ecosystem is 4% of total primary production, i.e. more similar to estimates for open ocean and coastal regions than for other upwelling or shelf systems averaging more than double this value. D 2002 Elsevier Science B.V. All rights reserved.

8 Resource and ecosystem variability, including regime shifts, in the Benguela Current System

Abstract Interannual and decadal-scale variability in abundance, distribution and biological characteristics are described for important living marine resources of the Benguela Current system including small pelagic fish, horse mackerel, hakes, snoek, rock lobster, Cape fur seals, Cape gannets and African penguins. Variability at the ecosystem level for the northern and southern subsystems is also described using trophodynamic indices that track structural changes in the ecosystem. Current understanding and analysis of observed variability in both resources and the ecosystem is reviewed, and the knowledge required for predicting resource and ecosystem variability and the causal factors that need to be considered are discussed. We highlight the need to improve understanding of the processes that are important in Benguela Current ecosystem, to identify what controls those processes, and to quantify such controls (particularly those acting on lower trophic levels) and the role of important species in the ecosystem. The kinds of predictions considered possible in the Benguela Current system are examined, and a series of steps is suggested to improve understanding of ecosystem and fisheries dynamics and to monitor key aspects of the ecosystem.

Multiple factors affecting South African anchovy recruitment in the spawning, transport and nursery areas

Despite high primary productivity, the yield of pelagic fish in the southern Benguela is relatively low compared to that in the Humboldt system. Part of the constraint may be the ability of pelagic fish to reproduce successfully in a strongly pulsed upwelling environment, where enrichment, retention and concentration mechanisms are less compatible than in Peru-Chile. Anchovy Engraulis capensis spawn upstream of the main upwelling centres on the food-poor, thermally stratified western Agulhas Bank, over a protracted summer season (October–February) when high wind speeds of 7–8 m˙s−1 are prevalent. Eggs spawned farther east, on the central or eastern Agulhas Bank, may be subject to increased cannibalism and advective losses, whereas those spawned farther west could be susceptible to heavy advective losses offshore during periods of strong southerly winds. Copepod concentrations are negatively correlated with spawner biomass on the western Bank and are inversely linked to high rates of gonad atresia in ancho...

Modelling stock dynamics in the southern Benguela ecosystem for the period 1978–2002

An ecosystem model of the southern Benguela was fitted to available time-series data for the period 1978–2002, to explore how changes in target fish populations in this ecosystem can be attributed to feeding interaction terms and population control patterns, the impact of fishing, and environmental forcing. Fishing patterns were estimated to explain only 2–3% of the variability in the time-series, whereas an estimated productivity forcing pattern applied to phytoplankton explained 4–12% of the variance represented by the sum of squares. Model settings describing prey vulnerability to their predators could explain around 40% of the variability in the time-series. Modelled stock dynamics in the southern Benguela ecosystem more closely represent observed timeseries when wasp-waist control by small pelagic fish is simulated. Overall, model simulations suggest that almost half the variance in the time-series can be explained based on a combination of fishing, vulnerability settings and productivity patterns. Variation in mortalities and prey preferences over time, as well as model fits in relation to available effort series, are discussed. The study advances a model with improved parameterization and credibility to assist with an ecosystem approach to South African fisheries management.

SIMULATIONS OF FISHING EFFECTS ON THE SOUTHERN BENGUELA FISH COMMUNITY USING AN INDIVIDUAL-BASED MODEL: LEARNING FROM A COMPARISON WITH ECOSIM

By applying an individual-based model (OSMOSE) to the southern Benguela ecosystem, a multispecies analysis is proposed, complementary to that provided by the application of ECOPATH/ECOSIM models. To reconstruct marine foodwebs, OSMOSE is based on the hypothesis that predation is a size-structured process. In all, 12 fish species, chosen for their importance in terms of biomass and catches, are explicitly modelled. Growth, reproduction and mortality parameters are required to model their dynamics and trophic interactions. Maps of mean spatial distribution of the species are compiled from published literature. Taking into account the spatial component is necessary because spatial co-occurrence determines potential interactions between predatory fish and prey fish of suitable size. To explore ecosystem effects of fishing, different fishing scenarios, previously examined using ECOSIM, are simulated using the OSMOSE model. They explore the effects of targeting fish species in the southern Benguela considered to be predators (Cape hake Merluccius capensis and M. paradoxus) or prey (anchovy Engraulis encrasicolus, sardine Sardinops sagax, round herring Etrumeus whiteheadi). Simulation results are compared and are generally consistent with those obtained using an ECOSIM model. This cross-validation appears to be a promising means of evaluating the robustness of model outputs, when separate validation of marine ecosystem models are still difficult to perform.

Comparing internal and external drivers in the southern Benguela and the southern and northern Humboldt upwelling ecosystems

Trophic models of three upwelling ecosystems, the southern Benguela (South African), southern Humboldt (Chilean) and northern Humboldt (Peruvian) systems, have been fitted to catch, abundance and fishing mortality time-series. Three drivers were considered during the model fitting: internal forcing by means of the trophic flow controls between the various interacting species groups, and two kinds of external forcing, namely fishing and the environment. The southern Benguela model was fitted to time-series data from 1978 to 2003, the southern Humboldt model to data from 1970 to 2004, and the northern Humboldt to data for a shorter period, 1995–2004. Fishing has been relatively carefully managed in the southern Benguela during the period modelled and previous studies found that most of the resource variability was attributed to internal trophodynamic forcing and to environmental forcing rather than to fishing. By comparison, fishing has been shown to have played a relatively major role in driving ecosystem changes observed in the southern and northern Humboldt models. Bearing in mind the different roles played by each of the drivers in these ecosystems, flow controls between interacting species groups, which improved the fits of the models, were compared across the three ecosystems to determine to what extent the three models supported the hypothesis that upwelling ecosystems function as wasp-waist systems. Secondly, environmental forcing was examined by searching for hypothetical forcing functions, affecting different levels of the foodweb, which improved the model fits. This was an attempt to start to uncover the processes that may be involved in linking the environment to observed ecosystem dynamics and changes in these upwelling ecosystems. Model results confirmed the important ecological role played by small pelagic fish in the studied upwelling ecosystems. For example, the fit of the southern Benguela model to time-series data of catch and abundance was similarly improved when anchovy/sardine–prey and anchovy/sardine–predator interactions were externally forced, supporting the wasp-waist hypothesis. In addition, although physical drivers and conditions may differ in their nature or merely their frequency and intensity between systems, and different fishing strategies operate in each of the three ecosystems, model results suggest that these effects are transferred through the ecosystems and manifest themselves as ecosystem changes and observed resource dynamics largely via interactions with small pelagic fish.