Modelling multi-species interactions in the Barents Sea ecosystem with special emphasis on minke whales and their interactions with cod, herring and capelin

The Barents Sea ecosystem, one of the most productive and commercially important ecosystems in the world, has experienced major fluctuations in species abundance the past five decades. Likely causes are natural variability, climate change, overfishing and predator–prey interactions. In this study, we use an age-length structured multi-species model (Gadget, Globally applicable Area-Disaggregated General Ecosystem Toolbox) to analyse the historic population dynamics of major fish and marine mammal species in the Barents Sea. The model was used to examine possible effects of a number of plausible biological and fisheries scenarios. The results suggest that changes in cod mortality from fishing or cod cannibalism levels have the largest effect on the ecosystem, while changes to the capelin fishery have had only minor effects. Alternate whale migration scenarios had only a moderate impact on the modelled ecosystem. Indirect effects are seen to be important, with cod fishing pressure, cod cannibalism and whale predation on cod having an indirect impact on capelin, emphasising the importance of multi-species modelling in understanding and managing ecosystems. Models such as the one presented here provide one step towards an ecosystem-based approach to fisheries management.     

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

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

Changes in the northern Gulf of St. Lawrence ecosystem estimated by inverse modelling: Evidence of a fishery-induced regime shift?

Mass-balance models have been constructed using inverse methodology for the northern Gulf of St. Lawrence for the mid-1980s, the mid-1990s, and the early 2000s to describe ecosystem structure, trophic group interactions, and the effects of fishing and predation on the ecosystem for each time period. Our analyses indicate that the ecosystem structure shifted dramatically from one previously dominated by demersal (cod, redfish) and small-bodied forage (e.g., capelin, mackerel, herring, shrimp) species to one now dominated by small-bodied forage species. Overfishing removed a functional group in the late 1980s, large piscivorous fish (primarily cod and redfish), which has not recovered 14 years after the cessation of heavy fishing. This has left only marine mammals as top predators during the mid-1990s, and marine mammals and small Greenland halibut during the early 2000s. Predation by marine mammals on fish increased from the mid-1980s to the early 2000s while predation by large fish on fish decreased. Capelin and shrimp, the main prey in each period, showed an increase in biomass over the three periods. A switch in the main predators of capelin from cod to marine mammals occurred, while Greenland halibut progressively replaced cod as shrimp predators. Overfishing influenced community structure directly through preferential removal of larger-bodied fishes and indirectly through predation release because larger-bodied fishes exerted top-down control upon other community species or competed with other species for the same prey. Our modelling estimates showed that a change in predation structure or flows at the top of the trophic system led to changes in predation at all lower trophic levels in the northern Gulf of St. Lawrence. These changes represent a case of fishery-induced regime shift.     

Abrupt transitions of the top-down controlled Black Sea pelagic ecosystem during 1960–2000: Evidence for regime-shifts under strong fishery exploitation and nutrient enrichment modulated by climate-induced variations

Functioning of the Black Sea ecosystem has profoundly changed since the early 1970s under cumulative effects of excessive nutrient enrichment, strong cooling/warming, over-exploitation of pelagic fish stocks, and population outbreak of gelatinous carnivores. Applying a set of criteria to the long-term (1960–2000) ecological time-series data, the present study demonstrates that the Black Sea ecosystem was reorganised during this transition phase in different forms of top-down controlled food web structure through successive regime-shifts of distinct ecological properties. The Secchi disc depth, oxic–anoxic interface zone, dissolved oxygen and hydrogen sulphide concentrations also exhibit abrupt transition between their alternate regimes, and indicate tight coupling between the lower trophic food web structure and the biogeochemical pump in terms of regime-shift events.The first shift, in 1973–1974, marks a switch from large predatory fish to small planktivore fish-controlled system, which persisted until 1989 in the form of increasing small pelagic and phytoplankton biomass and decreasing zooplankton biomass. The increase in phytoplankton biomass is further supported by a bottom-up contribution due to the cumulative response to high anthropogenic nutrient load and the concurrent shift of the physical system to the “cold climate regime” following its ∼20-year persistence in the “warm climate regime”. The end of the 1980s signifies the depletion of small planktivores and the transition to a gelatinous carnivore-controlled system. By the end of the 1990s, small planktivore populations take over control of the system again. Concomitantly, their top-down pressure when combined with diminishing anthropogenic nutrient load and more limited nutrient supply into the surface waters due to stabilizing effects of relatively warm winter conditions switched the “high production” regime of phytoplankton to its background “low production” regime.The Black Sea regime-shifts appear to be sporadic events forced by strong transient decadal perturbations, and therefore differ from the multi-decadal scale cyclical events observed in pelagic ocean ecosystems under low-frequency climatic forcing. The Black Sea observations illustrate that eutrophication and extreme fishery exploitation can indeed induce hysteresis in large marine ecosystems, when they can exert sufficiently strong forcing onto the system. They further illustrate the link between the disruption of the top predators, proliferation of new predator stocks, and regime-shift events. Examples of these features have been reported for some aquatic ecosystems, but are extremely limited for large marine ecosystems.     

Implications of variability on many time scales for scientific advice on sustainable management of living marine resources

The conceptual basis for understanding and management of living marine resources is built on three basic ecological principles developed in the first half of the past century: the law of the minimum, competitive exclusion, and succession. This paper highlights aspects of these principles that make them insufficient as a sound foundation for understanding and managing marine ecosystems, points out dangers of continuing to use approaches built on them, and presents alternatives which might be more appropriate and of lower risk.To do this, the paper considers variability of marine ecosystems on annual, medium and long-term time scales, highlighting that these scales correspond to less than, approximately equal to, and much greater than, the generation times of dominant predators in the systems. It also considers how each interval of variability may affect directly ecosystems which are controlled from the bottom up, top down, and middle outward, and how position and duration of forcing affect five types of responses: growth, maturation, recruitment, predation, and competition. Generally these five processes have manifestations at the scale of individuals, populations, and ecosystems, attention is drawn to which manifestations are the most significant for each duration and position of forcing.Effects of some combinations of duration of forcing and position of forcing can be explained reasonably well by conventional ecological theory. For other combinations, particularly forcing at time scales of predator generations on top-down or middle-out ecosystems, theory based on contest competition and equilibria are likely to be misleading. In these systems the major dynamics are transients, when many ecosystems are far from their carrying capacities, so scramble competition dominates, and the carrying capacity is not helpful in explaining the system dynamics. This review clarifies the sorts of questions that we should be asking, in order to begin to understand the transient behaviour of these non-equilibrium ecosystems. The answers to the new classes of questions may lead to great improvements in how ecosystems are managed, as well as how their variation is explained.     

Organisms as cooperative ecosystem engineers in intertidal flats

The importance of facilitative interactions and organismal ecosystem engineering for establishing the structure of communities is increasingly being recognised for many different ecosystems. For example, soft-bottom tidal flats host a wide range of ecosystem engineers, probably because the harsh physico-chemical environmental conditions render these species of particular importance for community structure and function. These environments are therefore interesting when focusing on how ecosystem engineers interact and the consequences of these interactions on community dynamics. In this review, we initially detail the influence on benthic systems of two kinds of ecosystem engineers that are particularly common in tidal flats. Firstly, we examine species providing biogenic structures, which are often the only source of habitat complexity in these environments. Secondly, we focus on species whose activities alter sediment stability, which is a crucial feature structuring the dynamics of communities in tidal flats. The impacts of these engineers on both environment and communities were assessed but in addition the interaction between ecosystem engineers was examined. Habitat cascades occur when one engineer favours the development of another, which in turn creates or modifies and improves habitat for other species. Non-hierarchical interactions have often been shown to display non-additive effects, so that the effects of the association cannot be predicted from the effects of individual organisms. Here we propose the term of “cooperative ecosystem engineering” when two species interact in a way which enhances habitat suitability as a result of a combined engineering effect. Finally, we conclude by describing the potential threats for ecosystem engineers in intertidal areas, potential effects on their interactions and their influence on communities and ecosystem function.     

Wasp-waist populations and marine ecosystem dynamics: Navigating the “predator pit” topographies

Many marine ecosystems exhibit a characteristic “wasp-waist” structure, where a single species, or at most several species, of small planktivorous fishes entirely dominate their trophic level. These species have complex life histories that result in radical variability that may propagate to both higher and lower trophic levels of the ecosystem. In addition, these populations have two key attributes: (1) they represent the lowest trophic level that is mobile, so they are capable of relocating their area of operation according to their own internal dynamics; (2) they may prey upon the early life stages of their predators, forming an unstable feedback loop in the trophic system that may, for example, precipitate abrupt regime shifts. Experience with the typical “boom-bust” dynamics of this type of population, and with populations that interact trophically with them, suggests a “predator pit” type of dynamics. This features a refuge from predation when abundance is very low, very destructive predation between an abundance level sufficient to attract interest from predators and an abundance level sufficient to satiate available predators, and, as abundance increases beyond this satiation point, decreasing specific predation mortality and population breakout. A simple formalism is developed to describe these dynamics. Examples of its application include (a) a hypothetical mechanism for progressive geographical habitat expansion at high biomass, (b) an explanation for the out-of-phase alternations of abundances of anchovies and sardines in many regional systems that appear to occur without substantial adverse interactions between the two species groups, and (c) an account of an interaction of environmental processes and fishery exploitation that caused a regime shift. The last is the example of the Baltic Sea, where the cod resource collapsed in concert with establishment of dominance of that ecosystem by the cod’s ‘wasp-waist” prey, herring and sprat.     

Ecosystem exploitation and trophodynamic indicators: A comparison between the Northern Adriatic Sea and Southern New England

In an ecosystem-based resource management context, it is crucial to assess the relationships between community structure and ecosystem function and how those relationships change with resource extraction. To elucidate how changes in resource use can affect community structure and ecosystem function, we executed a comparative analysis of two different ecosystems subjected to notable fishing pressure. We contrasted the Northern Adriatic Sea (NAS) and Southern New England (SNE) ecosystems by examining outputs from comparable steady-state models. Both ecosystems have relatively high fishing pressure and a high biomass of benthic invertebrates. The basic structure of the food webs shows differences both in the number and definition of the functional groups, as described in the models. Fisheries, on the contrary, show similarities both in terms of catches and discards. Almost all statistics summarizing the structure and flows showed values three times higher in the SNE than in the NAS ecosystem, but despite this difference the two ecosystems exhibited similar, overall properties. Biomass ratios and the Mixed Trophic Impact (MTI) analysis showed that both ecosystems are dominated by the benthic compartment. Removing the biomass effect, however, shows a clear top-down effect, with a high rank achieved by fishing activities. In general terms, the low mean trophic level of catches and the high primary production required (PPR) values result in a high overexploitation level of the ecosystem, as highlighted by the L index. We conclude by exploring how comparative studies will continue to be valuable as ecosystem-based management is further implemented.     

Intraguild predation and partial consumption of blue sharks Prionace glauca by Cape fur seals Arctocephalus pusillus pusillus

The top-down effects of predators on ecosystem structure and dynamics have been studied increasingly. However, the nature and consequence of trophic interactions between upper-trophic-level predators have received considerably less attention. This is especially the case in marine systems due to the inherent challenges of studying highly mobile marine species. Here we describe the first documentation of asymmetrical intraguild predation by a pinniped predator on a mid-sized predatory shark. The report is based on direct observations in South African waters, in which free-swimming blue sharks Prionace glauca were captured and partially consumed by Cape fur seals Arctocephalus pusillus pusillus. These observations are important not just for understanding the interactions between these two species but more broadly for their implications in understanding the trophic ecology of pinnipeds, many populations of which have increased while numerous shark populations have declined.     

Review of indicators in fisheries management – a development perspective

A review of indicators for an ecosystem approach to fisheries management is presented, focusing on multispecies fisheries and limited resources for assessments and implementation, as often is the case in developing countries. Emphasizing the need to link indicators to management objectives, indicators from the literature are grouped into four categories, relating to the immediate fisheries resource base (single-species and multispecies indicators) and the wider ecosystem (habitat structure and ecosystem functioning). The usefulness of these indicators is assessed along three dimensions of acceptability among stakeholders, observability, and relation to fisheries management using a traffic light approach. The top ranking indicators are highlighted as a generally good start for any particular fishery management case. It is, however, argued that, even with similar management objectives, indicators need to be specific to both ecosystems and the institutional set-up if they are to be effective for management, and that indicators may consequently differ considerably between individual management applications.     

Ecosystem services of the tropical seascape: interactions,substitutions and restoration

The tropical coastal “seascape” often includes a patchwork of mangroves, seagrass beds, and coral reefs that produces a variety of natural resources and ecosystem services. By looking into a limited number of attempts at substitution and restoration of ecosystem services (e.g. artificial reefs, aquaculture in mangroves, artificial seawalls), we address the questions: (1) To what degree can technologies substitute for ecosystem services in the seascape? (2) How can ecosystem restoration reestablish not only the functions of direct value to humans, but also the ability of the systems to cope with future disturbance? Substitutions often imply the replacement of a function provided free by a solar powered, self-repairing resilient ecosystem, with a fossil-fuel-powered, expensive, artificial substitute that needs maintenance. Further, restoration usually does not focus on large-scale processes such as the physical, biological and biogeochemical interactions between mangroves, seagrass beds and coral reefs. Nonetheless, restoration might be the only viable management alternative when the system is essentially locked into an undesired community state (stability domain) after a phase-shift. We conclude that ecosystem services cannot be readily replaced, restored or sustained without extensive knowledge of the dynamics, multifunctionality and interconnectedness of ecosystems.     

南海典型珊瑚礁生态系统健康评价方法研究

本研究基于系统复杂性原理, 提出了南海典型珊瑚礁生态系统健康评价方法——组织力及系统功能评价法。该方法的评价框架包括珊瑚礁生态系统结构及功能、珊瑚礁生态系统与其他海洋生态系统间关系、与人类社会生态系统间关系、珊瑚礁生态系统发展制约因素等四大模块; 评价指标体系分为目标层、准则层、因素层、要素层等6个层级共计67个指标。同时, 本研究系统地提出了珊瑚礁生态系统健康评价的流程, 提高了珊瑚礁健康评价工作的规范性和时效性。应用组织力及系统功能评价法对2011—2018年西沙群岛珊瑚礁生态系统进行了健康评价, 结果显示自2011年起健康指数年均下降2.3%~2.4%, 除2012年、2015年健康指数稍有回升外, 其余年份均呈下降走势, 整体评价为亚健康状态。根据七连屿周边岛屿的实地考察和居民生活问卷调查结果, 2006—2008年的渔业发展、2012年的海星泛滥和2014—2015年的岛礁建设均对珊瑚礁生态系统造成了较大影响, 使10年内渔业产量下降了50%~80%。这与应用组织力及系统功能评价法的评价结果相吻合, 侧面验证了该评价方法的科学性和可行性。     

Recent climate forcing and physical oceanographic changes in Northern Hemisphere regions: A review and comparison of four marine ecosystems

As part of a project comparing the structure and function of four marine ecosystems off Norway and the United States, this paper examines the oceanographic responses to climate forcing, with emphasis on recent changes. The four Northern Hemisphere ecosystems include two in the Pacific Ocean (Bering Sea and Gulf of Alaska) and two in the Atlantic Ocean (Georges Bank/Gulf of Maine and the Barents/Norwegian Seas). Air temperatures, wind forcing and heat fluxes over the four regions are compared as well as ocean hydrography and sea-ice conditions where seasonal sea ice is found. The long-term interannual variability in air temperatures, winds and net heat fluxes show strong similarity between adjacent ecosystems and within subregions of an ecosystem, but no significant correlations between Pacific and Atlantic ecosystems and few across the Atlantic. In spite of the lack of correlation between climate forcing and ocean conditions between most of the ecosystems, recent years have seen record or near record highs in air and sea temperatures in all ecosystems. The apparent causes of the warming differ. In the Atlantic, they appear to be due to advection, while in the Pacific temperatures are more closely linked to air-sea heat exchanges. Advection is also responsible for the observed changes in salinity in the Atlantic ecosystems (generally increasing salinity in the Barents and Norwegian Seas and decreasing in the Gulf of Maine and Georges Bank) while salinity changes in the Gulf of Alaska are largely related to increased local runoff.     

Three-dimensional mapping of red stingray (Dasyatis akajei) movement with reference to bottom topography

Most demersal fishes maintain strong relations with bottom substrates and bottom depths and/or topography during their lives. It is important to know these relations to for understand their lives. In Tokyo Bay, red stingray, Dasyatis akajei, classified as near-threatened species by IUCN, has increased since the 1980s. It is a top predator and engages in ecosystem engineer by mixing the sand bed surface through burring behavior, and greatly influences a coastal ecosystem. It is reported that this species invades in plage and tidal flats and has sometimes injured beachgoers and people gathering clams in Tokyo bay. Thus, it is necessary to know its behavior and habitat use to avoid accidents and to better conserve the biodiversity of ecosystems. However, previous studies have not examined its relationship with the bottom environment. This study aims to describe its behavior in relation to the bottom environment. We sounded three dimensional bottom topography of their habitat off Kaneda Cove in Tokyo Bay with interferometric sidescan sonar system and traced the movement of red stingrays by attaching a data logger system to survey their migration. The results revealed that red stingray repeated vertical movement between the surface and bottom, and used not only sand beds but also rocky beds.

     

Ecosystem structure and resilience—A comparison between the Norwegian and the Barents Sea

Abundance and biomass of the most important fish species inhabited the Barents and Norwegian Sea ecosystems have shown considerable fluctuations over the last decades. These fluctuations connected with fishing pressure resulted in the trophic structure alterations of the ecosystems. Resilience and other theoretical concepts (top-down, wasp-waste and bottom-up control, trophic cascades) were viewed to examine different response of the Norwegian and Barents Sea ecosystems on disturbing forces. Differences in the trophic structure and functioning of Barents and Norwegian Sea ecosystems as well as factors that might influence the resilience of the marine ecosystems, including climatic fluctuation, variations in prey and predator species abundance, alterations in their regular migrations, and fishing exploitation were also considered. The trophic chain lengths in the deep Norwegian Sea are shorter, and energy transfer occurs mainly through the pelagic fish/invertebrates communities. The shallow Barents Sea is characterized by longer trophic chains, providing more energy flow into their benthic assemblages. The trophic mechanisms observed in the Norwegian Sea food webs dominated by the top-down control, i.e. the past removal of Norwegian Spring spawning followed by zooplankton development and intrusion of blue whiting and mackerel into the area. The wasp-waist response is shown to be the most pronounced effect in the Barents Sea, related to the position of capelin in the ecosystem; large fluctuations in the capelin abundance have been strengthened by intensive fishery. Closer links between ecological and fisheries sciences are needed to elaborate and test various food webs and multispecies models available.     

Challenging paradigms in estuarine ecology and management

For many years, estuarine science has been the ‘poor relation’ in aquatic research - freshwater scientists ignored estuaries as they tended to get confused by salt and tides, and marine scientists were more preoccupied by large open systems. Estuaries were merely regarded by each group as either river mouths or sea inlets respectively. For the past four decades, however, estuaries (and other transitional waters) have been regarded as being ecosystems in their own right. Although often not termed as such, this has led to paradigms being generated to summarise estuarine structure and functioning and which relate to both the natural science and management of these systems. This paper defines, details and affirms these paradigms that can be grouped into those covering firstly the science (definitions, scales, linkages, productivity, tolerances and variability) and secondly the management (pressures, valuation, health and services) of estuaries. The more ‘science’ orientated paradigms incorporate the development and types of ecotones, the nature of stressed and variable systems (with specific reference to resilience and redundancy), the relationship between generalists and specialists produced by environmental tolerance, the relevance of scale in relation to functioning and connectivity, the sources of production and degree of productivity, the biodiversity-ecosystem functioning and the stress-subsidy debates. The more ‘management’ targeted paradigms include the development and effects of exogenic unmanaged pressures and endogenic managed pressures, the perception of health and the ability to manage estuaries (related to internal and external influences), and the influence of all of these on the production of ecosystem services and societal benefits.     

Residency,site fidelity and habitat use of Atlantic cod (Gadus morhua) at an offshore wind farm using acoustic telemetry

Because offshore wind energy development is fast growing in Europe it is important to investigate the changes in the marine environment and how these may influence local biodiversity and ecosystem functioning. One of the species affected by these ecosystem changes is Atlantic cod (Gadus morhua), a heavily exploited, commercially important fish species. In this research we investigated the residency, site fidelity and habitat use of Atlantic cod on a temporal scale at windmill artificial reefs in the Belgian part of the North Sea. Acoustic telemetry was used and the Vemco VR2W position system was deployed to quantify the movement behaviour. In total, 22 Atlantic cod were tagged and monitored for up to one year. Many fish were present near the artificial reefs during summer and autumn, and demonstrated strong residency and high individual detection rates. When present within the study area, Atlantic cod also showed distinct habitat selectivity. We identified aggregation near the artificial hard substrates of the wind turbines. In addition, a clear seasonal pattern in presence was observed. The high number of fish present in summer and autumn alternated with a period of very low densities during the winter period.