A regime shift in the Southeast Greenland marine ecosystem.

Two major oceanographic changes have recently propagated through several trophic levels in coastal areas of Southeast Greenland (SEG). Firstly, the amount of drift-ice exported from the Fram Strait and transported with the East Greenland Current (EGC) has decreased significantly over the past two decades, and a main tipping element (summer sea ice) has virtually disappeared since 2003 leading to a regime shift in oceanographic and ecological conditions in the region. The following 20-year period with low or no coastal sea ice is unique in the 200-year history of ice observations in the region, and the regime shift is also obvious in the volume of ice export through the Fram Strait after 2013. In the same period, the temperature of the EGC south of 73.5 N has increased significantly (>2°C) since 1980. Secondly, the warm Irminger Current, which advects warm, saline Atlantic Water into the region, has become warmer since 1990. The lack of pack ice in summer together with a warming ocean generated cascading effects on the ecosystem in SEG that are manifested in a changed fish fauna with an influx of boreal species in the south and the subarctic capelin further north. At higher trophic levels there has been an increase in the abundance of several boreal cetaceans (humpback, fin, killer, and pilot whales and dolphins) that are either new to this area or occur in historically large numbers. It is estimated that the new cetacean species in SEG are responsible for an annual predation level of 700,000 tons of fish. In addition, predation on krill species is estimated at >1,500,000 tons mainly consumed by fin whales. Simultaneously, there has been a reduction in the abundance and catches of narwhals and walruses in SEG and it is suggested that these species have been impacted by the habitat changes.

Importance of the Lunar Cycle on Mesopelagic Foraging by Atlantic Bluefin Tuna in the Upwelling Area of the Strait of Messina (Central Mediterranean Sea).

The influence of the lunar cycle on bluefin tuna foraging in the upwelling area of the Strait of Messina was investigated by exploring trophic interaction with mesopelagic fish and cephalopod prey. To focus on how the lunar cycle could affect availability of mesopelagic prey for this predator, we tested potential differences in the diet related to each lunar phase. Moreover, we considered two potential impacts of the lunar cycle: the lunar irradiance and the strength of currents. Overall, 2672 prey items were mesopelagic fish and cephalopods, representing 60.7% of overall diet by number. The main mesopelagic fish prey items were lanternfishes and dragonfishes, while Onychoteuthis banksii was the most important cephalopod prey. In summary, the Strait of Messina has highly specific hydrodynamic and biological features which strongly depend on upwelling currents, which in turn are influenced by the lunar cycle (new and full moon with strong currents, quarters with fewer currents). Upwelling causes water mixing, bringing to the surface a large amount of mesopelagic fauna which become more readily available to tuna. Lunar irradiance contributes to the variation of prey composition, increasing the success of visual predation on mesopelagic resources at high light in the water column.

Economic expenditures by recreational anglers in a recovering atlantic bluefin tuna fishery.

The recent return of Atlantic bluefin tuna to northern Europe following the recovery of the east Atlantic stock has sparked substantial public and scientific interest. This is particularly true for recreational anglers in Denmark, who often consider Atlantic bluefin tuna to be the catch of a lifetime. This attitude has previously sustained a substantial recreational fishery for bluefin tuna with annual tournaments in Denmark, which peaked in the 1950s before the subsequent collapse of the stock during the 1960s. Several scientific tagging programs have recruited recreational anglers in recent years to help catch and release tagged bluefin tuna. The anglers' investment of time and money in the scientific tagging projects indicate that the recreational fishery could recover in the future. However, the economic aspects of a potential future recreational bluefin tuna fishery remain unknown. We surveyed anglers participating in a scientific catch and release bluefin tuna fishery in Denmark across three years (2018-2020) and calculated the total annual expenditures associated with the activities. Additionally, we estimated the magnitude of the negative impact (i.e., incidental mortalities) on the bluefin tuna stock. Our results show that total annual expenditures by the recreational anglers approached 1,439,540€, totaling 4,318,620€ between 2018 and 2020. We found that recreational bluefin tuna anglers had mean annual expenditures directly related to the bluefin tuna fishing between 7,047€ and 2,176€ with an associated mortality impact on the stock of less than 1 tonne annually. By comparing the mortality impact to the expenditures, we estimate that each dead Atlantic bluefin tuna during the three study years generated 398,163€ in mean annual expenditures, equivalent to approximately 1636€ kg-1. Our study demonstrates significant economic expenditures among recreational anglers who target Atlantic bluefin tuna. This provides a clear example of how a recovery of marine natural capital and related ecosystem services can support development in the blue economy.

First tagging data on large Atlantic bluefin tuna returning to Nordic waters suggest repeated behaviour and skipped spawning.

Atlantic bluefin tuna (Thunnus thynnus; ABFT) is one of the most iconic fish species in the world. Recently, after being very rare for more than half a century, large bluefin tunas have returned to Nordic waters in late summer and autumn, marking the return of the largest predatory fish in Nordic waters. By tagging 18 bluefin tunas with electronic tags (pop-up satellite archival tags), we show that bluefin tuna observed in Nordic waters undertake different migration routes, with individuals migrating into the western Atlantic Ocean, while others stay exclusively in the eastern Atlantic and enter the Mediterranean Sea to spawn. We additionally present evidence of possible skipped spawning inferred from behavioural analyses. In Nordic waters, ABFT are primarily using the upper water column, likely reflecting feeding activity. The results support the hypothesis that ABFT migrating to Nordic waters return to the same general feeding area within the region on an annual basis. These observations may have important implications for management because (1) tunas that come into Nordic waters might represent only a few year classes (as evidenced by a narrow size range), and thus may be particularly vulnerable to area-specific exploitation, and (2) challenge the assumption of consecutive spawning in adult Atlantic bluefin tuna, as used in current stock assessment models. Without careful management and limited exploitation of this part of the ABFT population, the species' return to Nordic waters could be short-lived.

Temperature-dependent adaptation allows fish to meet their food across their species' range.

In seasonal environments, timing is everything: Ecosystem dynamics are controlled by how well predators can match their prey in space and time. This match of predator and prey is thought to be particularly critical for the vulnerable larval life stages of many fish, where limited parental investment means that population survival can depend on how well larvae match the timing of their food. We develop and apply novel metrics of thermal time to estimate the timing of unobserved stages of fish larvae and their prey across the north Atlantic. The result shows that previously identified life-history strategies are adaptive in that they allow parents to "predict" a beneficial environment for their offspring and meet larval fish food timing that varies by 99 days across a species' range.

The Baltic Sea as a time machine for the future coastal ocean.

Coastal global oceans are expected to undergo drastic changes driven by climate change and increasing anthropogenic pressures in coming decades. Predicting specific future conditions and assessing the best management strategies to maintain ecosystem integrity and sustainable resource use are difficult, because of multiple interacting pressures, uncertain projections, and a lack of test cases for management. We argue that the Baltic Sea can serve as a time machine to study consequences and mitigation of future coastal perturbations, due to its unique combination of an early history of multistressor disturbance and ecosystem deterioration and early implementation of cross-border environmental management to address these problems. The Baltic Sea also stands out in providing a strong scientific foundation and accessibility to long-term data series that provide a unique opportunity to assess the efficacy of management actions to address the breakdown of ecosystem functions. Trend reversals such as the return of top predators, recovering fish stocks, and reduced input of nutrient and harmful substances could be achieved only by implementing an international, cooperative governance structure transcending its complex multistate policy setting, with integrated management of watershed and sea. The Baltic Sea also demonstrates how rapidly progressing global pressures, particularly warming of Baltic waters and the surrounding catchment area, can offset the efficacy of current management approaches. This situation calls for management that is (i) conservative to provide a buffer against regionally unmanageable global perturbations, (ii) adaptive to react to new management challenges, and, ultimately, (iii) multisectorial and integrative to address conflicts associated with economic trade-offs.

A global mismatch in the protection of multiple marine biodiversity components and ecosystem services.

The global loss of biodiversity threatens unique biota and the functioning and services of ecosystems essential for human wellbeing. To safeguard biodiversity and ecosystem services, designating protected areas is crucial; yet the extent to which the existing placement of protection is aligned to meet these conservation priorities is questionable, especially in the oceans. Here we investigate and compare global patterns of multiple biodiversity components (taxonomic, phylogenetic and functional), ecosystem services and human impacts, with the coverage of marine protected areas across a nested spatial scale. We demonstrate a pronounced spatial mismatch between the existing degree of protection and all the conservation priorities above, highlighting that neither the world's most diverse, nor the most productive ecosystems are currently the most protected ecosystems. Furthermore, we show that global patterns of biodiversity, ecosystem services and human impacts are poorly correlated, hence complicating the identification of generally applicable spatial prioritization schemes. However, a hypothetical "consensus approach" would have been able to address all these conservation priorities far more effectively than the existing degree of protection, which at best is only marginally better than a random expectation. Therefore, a holistic perspective is needed when designating an appropriate degree of protection of marine conservation priorities worldwide.

Global patterns in marine predatory fish.

Large teleost (bony) fish are a dominant group of predators in the oceans and constitute a major source of food and livelihood for humans. These species differ markedly in morphology and feeding habits across oceanic regions; large pelagic species such as tunas and billfish typically occur in the tropics, whereas demersal species of gadoids and flatfish dominate boreal and temperate regions. Despite their importance for fisheries and the structuring of marine ecosystems, the underlying factors determining the global distribution and productivity of these two groups of teleost predators are poorly known. Here, we show how latitudinal differences in predatory fish can essentially be explained by the inflow of energy at the base of the pelagic and benthic food chain. A low productive benthic energy pathway favours large pelagic species, whereas equal productivities support large demersal generalists that outcompete the pelagic specialists. Our findings demonstrate the vulnerability of large teleost predators to ecosystem-wide changes in energy flows and hence provide key insight to predict the responses of these important marine resources under global change.

Ocean warming expands habitat of a rich natural resource and benefits a national economy.

Geographic redistribution of living natural resources changes access and thereby harvesting opportunities between countries. Internationally shared fish resources can be sensitive to shifts in the marine environment and this may have great impact on the economies of countries and regions that rely most heavily on fisheries to provide employment and food supply. Here we present a climate change-related biotic expansion of a rich natural resource with substantial economic consequences, namely the appearance of northeast Atlantic mackerel (Scomber scombrus) in Greenlandic waters. In recent years, the summer temperature has reached record highs in the Irminger Current, and this development has expanded the available and realized mackerel habitat in time and space. Observations in the Irminger Current in east Greenland in 2011 of this temperature-sensitive epipelagic fish were the first records so far northwest in the Atlantic. This change in migration pattern was followed by a rapid development of a large-scale fishery of substantial importance for the national economy of Greenland (23% of Greenland's export value of all goods in 2014). A pelagic trawl survey was conducted in mid-summer 2014 and the results showed that the bulk of ~1 million Mg (=t) of mackerel in the Irminger Current in southeast Greenland were located in the relatively warm (>8.5°C) surface layer. Mackerel was also observed in southwest Greenland. Finally, 15 CMIP5 Earth System Model projections of future marine climate were used to evaluate the epipelagic environment in Greenland. These projections for moderate and high CO2 emission scenarios (representative concentration pathways [RCP] 4.5 and 8.5) suggest how the available mackerel habitat may expand further in space and time. Overall, our results indicate that, if the stock remains large, productive, and continues its current migration pattern, then climate change has provided Greenland with a new unique opportunity for commercial exploitation. However, positive cases like this should not be cherry-picked and misused as arguments against timely and effective mitigation of climate change.

Has eutrophication promoted forage fish production in the Baltic Sea?

Reducing anthropogenic nutrient inputs is a major policy goal for restoring good environmental status of coastal marine ecosystems. However, it is unclear to what extent reducing nutrients would also lower fish production and fisheries yields. Empirical examples of changes in nutrient loads and concurrent fish production can provide useful insights to this question. In this paper, we investigate to what extent a multi-fold increase in nutrient loads from the 1950s to 1980s enhanced forage fish production in the Baltic Sea. We use monitoring data on fish stock dynamics covering the period of the nutrient increase, combined with nutrient concentrations from a 3-dimensional coupled physical-biogeochemical ocean model. The results suggest that nutrient enrichment enhanced the biomass level of forage fish by up to 50 % in some years and areas due to increased body weight of fish. However, the trends in fish biomasses were generally decoupled from changes in nutrient concentrations.

A cascade of warming impacts brings bluefin tuna to Greenland waters.

Rising ocean temperatures are causing marine fish species to shift spatial distributions and ranges, and are altering predator-prey dynamics in food webs. Most documented cases of species shifts so far involve relatively small species at lower trophic levels, and consider individual species in ecological isolation from others. Here, we show that a large highly migratory top predator fish species has entered a high latitude subpolar area beyond its usual range. Bluefin tuna, Thunnus thynnus Linnaeus 1758, were captured in waters east of Greenland (65°N) in August 2012 during exploratory fishing for Atlantic mackerel, Scomber scombrus Linnaeus 1758. The bluefin tuna were captured in a single net-haul in 9-11 °C water together with 6 tonnes of mackerel, which is a preferred prey species and itself a new immigrant to the area. Regional temperatures in August 2012 were historically high and contributed to a warming trend since 1985, when temperatures began to rise. The presence of bluefin tuna in this region is likely due to a combination of warm temperatures that are physiologically more tolerable and immigration of an important prey species to the region. We conclude that a cascade of climate change impacts is restructuring the food web in east Greenland waters.

Fishing out collective memory of migratory schools.

Animals form groups for many reasons, but there are costs and benefits associated with group formation. One of the benefits is collective memory. In groups on the move, social interactions play a crucial role in the cohesion and the ability to make consensus decisions. When migrating from spawning to feeding areas, fish schools need to retain a collective memory of the destination site over thousands of kilometres, and changes in group formation or individual preference can produce sudden changes in migration pathways. We propose a modelling framework, based on stochastic adaptive networks, that can reproduce this collective behaviour. We assume that three factors control group formation and school migration behaviour: the intensity of social interaction, the relative number of informed individuals and the strength of preference that informed individuals have for a particular migration area. We treat these factors independently and relate the individuals' preferences to the experience and memory for certain migration sites. We demonstrate that removal of knowledgeable individuals or alteration of individual preference can produce rapid changes in group formation and collective behaviour. For example, intensive fishing targeting the migratory species and also their preferred prey can reduce both terms to a point at which migration to the destination sites is suddenly stopped. The conceptual approaches represented by our modelling framework may therefore be able to explain large-scale changes in fish migration and spatial distribution.

Combined effects of global climate change and regional ecosystem drivers on an exploited marine food web.

Changes in climate, in combination with intensive exploitation of marine resources, have caused large-scale reorganizations in many of the world's marine ecosystems during the past decades. The Baltic Sea in Northern Europe is one of the systems most affected. In addition to being exposed to persistent eutrophication, intensive fishing, and one of the world's fastest rates of warming in the last two decades of the 20th century, accelerated climate change including atmospheric warming and changes in precipitation is projected for this region during the 21st century. Here, we used a new multimodel approach to project how the interaction of climate, nutrient loads, and cod fishing may affect the future of the open Central Baltic Sea food web. Regionally downscaled global climate scenarios were, in combination with three nutrient load scenarios, used to drive an ensemble of three regional biogeochemical models (BGMs). An Ecopath with Ecosim food web model was then forced with the BGM results from different nutrient-climate scenarios in combination with two different cod fishing scenarios. The results showed that regional management is likely to play a major role in determining the future of the Baltic Sea ecosystem. By the end of the 21st century, for example, the combination of intensive cod fishing and high nutrient loads projected a strongly eutrophicated and sprat-dominated ecosystem, whereas low cod fishing in combination with low nutrient loads resulted in a cod-dominated ecosystem with eutrophication levels close to present. Also, nonlinearities were observed in the sensitivity of different trophic groups to nutrient loads or fishing depending on the combination of the two. Finally, many climate variables and species biomasses were projected to levels unseen in the past. Hence, the risk for ecological surprises needs to be addressed, particularly when the results are discussed in the ecosystem-based management context.

Spawning of bluefin tuna in the Black Sea: historical evidence, environmental constraints and population plasticity.

The lucrative and highly migratory Atlantic bluefin tuna, Thunnus thynnus (Linnaeus 1758; Scombridae), used to be distributed widely throughout the north Atlantic Ocean, Mediterranean Sea and Black Sea. Its migrations have supported sustainable fisheries and impacted local cultures since antiquity, but its biogeographic range has contracted since the 1950s. Most recently, the species disappeared from the Black Sea in the late 1980s and has not yet recovered. Reasons for the Black Sea disappearance, and the species-wide range contraction, are unclear. However bluefin tuna formerly foraged and possibly spawned in the Black Sea. Loss of a locally-reproducing population would represent a decline in population richness, and an increase in species vulnerability to perturbations such as exploitation and environmental change. Here we identify the main genetic and phenotypic adaptations that the population must have (had) in order to reproduce successfully in the specific hydrographic (estuarine) conditions of the Black Sea. By comparing hydrographic conditions in spawning areas of the three species of bluefin tunas, and applying a mechanistic model of egg buoyancy and sinking rate, we show that reproduction in the Black Sea must have required specific adaptations of egg buoyancy, fertilisation and development for reproductive success. Such adaptations by local populations of marine fish species spawning in estuarine areas are common as is evident from a meta-analysis of egg buoyancy data from 16 species of fish. We conclude that these adaptations would have been necessary for successful local reproduction by bluefin tuna in the Black Sea, and that a locally-adapted reproducing population may have disappeared. Recovery of bluefin tuna in the Black Sea, either for spawning or foraging, will occur fastest if any remaining locally adapted individuals are allowed to survive, and by conservation and recovery of depleted Mediterranean populations which could through time re-establish local Black Sea spawning and foraging.

Impact of climate change on fish population dynamics in the Baltic sea: a dynamical downscaling investigation.

Understanding how climate change, exploitation and eutrophication will affect populations and ecosystems of the Baltic Sea can be facilitated with models which realistically combine these forcings into common frameworks. Here, we evaluate sensitivity of fish recruitment and population dynamics to past and future environmental forcings provided by three ocean-biogeochemical models of the Baltic Sea. Modeled temperature explained nearly as much variability in reproductive success of sprat (Sprattus sprattus; Clupeidae) as measured temperatures during 1973-2005, and both the spawner biomass and the temperature have influenced recruitment for at least 50 years. The three Baltic Sea models estimate relatively similar developments (increases) in biomass and fishery yield during twenty-first century climate change (ca. 28 % range among models). However, this uncertainty is exceeded by the one associated with the fish population model, and by the source of global climate data used by regional models. Knowledge of processes and biases could reduce these uncertainties.

Stable isotope evidence for late medieval (14th-15th C) origins of the eastern Baltic cod (Gadus morhua) fishery.

Although recent historical ecology studies have extended quantitative knowledge of eastern Baltic cod (Gadus morhua) exploitation back as far as the 16th century, the historical origin of the modern fishery remains obscure. Widespread archaeological evidence for cod consumption around the eastern Baltic littoral emerges around the 13th century, three centuries before systematic documentation, but it is not clear whether this represents (1) development of a substantial eastern Baltic cod fishery, or (2) large-scale importation of preserved cod from elsewhere. To distinguish between these hypotheses we use stable carbon and nitrogen isotope analysis to determine likely catch regions of 74 cod vertebrae and cleithra from 19 Baltic archaeological sites dated from the 8th to the 16th centuries. δ(13)C and δ(15)N signatures for six possible catch regions were established using a larger sample of archaeological cod cranial bones (n = 249). The data strongly support the second hypothesis, revealing widespread importation of cod during the 13th to 14th centuries, most of it probably from Arctic Norway. By the 15th century, however, eastern Baltic cod dominate within our sample, indicating the development of a substantial late medieval fishery. Potential human impact on cod stocks in the eastern Baltic must thus be taken into account for at least the last 600 years.

Could seals prevent cod recovery in the Baltic Sea?

Fish populations are increasingly affected by multiple human and natural impacts including exploitation, eutrophication, habitat alteration and climate change. As a result many collapsed populations may have to recover in ecosystems whose structure and functioning differ from those in which they were formerly productive and supported sustainable fisheries. Here we investigate how a cod (Gadus morhua) population in the Baltic Sea whose biomass was reduced due to a combination of high exploitation and deteriorating environmental conditions might recover and develop in the 21st century in an ecosystem that likely will change due to both the already started recovery of a cod predator, the grey seal Halichoerus grypus, and projected climate impacts. Simulation modelling, assuming increased seal predation, fishing levels consistent with management plan targets and stable salinity, shows that the cod population could reach high levels well above the long-term average. Scenarios with similar seal and fishing levels but with 15% lower salinity suggest that the Baltic will still be able to support a cod population which can sustain a fishery, but biomass and yields will be lower. At present knowledge of cod and seal interactions, seal predation was found to have much lower impact on cod recovery, compared to the effects of exploitation and salinity. These results suggest that dual management objectives (recovery of both seal and cod populations) are realistic but success in achieving these goals will also depend on how climate change affects cod recruitment.

Multi-decadal responses of a cod (Gadus morhua) population to human-induced trophic changes, fishing, and climate.

Understanding how human impacts have interacted with natural variability to affect populations and ecosystems is required for sustainable management and conservation. The Baltic Sea is one of the few large marine ecosystems worldwide where the relative contribution of several key forcings to changes in fish populations can be analyzed with empirical data. In this study we investigate how climate variability and multiple human impacts (fishing, marine mammal hunting, eutrophication) have affected multi-decadal scale dynamics of cod in the Baltic Sea during the 20th century. We document significant climate-driven variations in cod recruitment production at multi-annual timescales, which had major impacts on population dynamics and the yields to commercial fisheries. We also quantify the roles of marine mammal predation, eutrophication, and exploitation on the development of the cod population using simulation analyses, and show how the intensity of these forcings differed over time. In the early decades of the 20th century, marine mammal predation and nutrient availability were the main limiting factors; exploitation of cod was still relatively low. During the 1940s and subsequent decades, exploitation increased and became a dominant forcing on the population. Eutrophication had a relatively minor positive influence on cod biomass until the 1980s. The largest increase in cod biomass occurred during the late 1970s, following a long period of hydrographically related above-average cod productivity coupled to a temporary reduction in fishing pressure. The Baltic cod example demonstrates how combinations of different forcings can have synergistic effects and consequently dramatic impacts on population dynamics. Our results highlight the potential and limitations of human manipulations to influence predator species and show that sustainable management can only be achieved by considering both anthropogenic and naturally varying processes in a common framework.