The dangers of ignoring stock complexity in fishery management: the case of the North Sea cod

The plight of the marine fisheries is attracting increasing attention as unsustainably high exploitation levels, exacerbated by more extreme climatic conditions, are driving stocks to the point of collapse. The North Atlantic cod (Gadus morhua), a species which until recently formed a major component of the demersal fisheries, has undergone significant declines across its range. The North Sea stock is typical of many, with a spawning stock biomass that has remained below the safe biological limit since 2000 and recruitment levels near the lowest on record. Cod within the North Sea are currently managed as a single stock, and yet mounting empirical evidence supports the existence of a metapopulation of regionally variable, genetically distinct, sub-stocks. Applying the same management strategies to multiple stocks that differ in their resilience to exploitation inevitably results in the overfishing and likely collapse of the weaker components. Indeed, recent studies have identified two North Sea spawning stocks that have undergone disproportionally large collapses with very substantial reductions in egg production. Similarly affected cod stocks in the northwest Atlantic have shown little evidence of recovery, despite fishery closures. The possible implications of ignoring sub-structuring within management units for biocomplexity, local adaptation and ecosystem stability are considered.     

DNA-based identification of larval cod in stomach contents of predatory fishes

Predator-prey interactions play an influential role in determining the demographics of a population or species. In the Northwest Atlantic, Atlantic cod, Gadus morhua, once the basis of a lucrative commercial fishery, have not recovered despite regulations imposed on the fishery to reduce harvest rates. One possible reason for the lack of recovery is that high predation pressure on juvenile and larval stages, particularly from species such as herring and mackerel, may regulate the abundance of cod. However, traditional methods used to identify larval cod and haddock often fail when applied to partially digested remains. Here, we described a DNA-based assay to identify the presence of digested cod remains from the stomachs of predatory fish species. After development, the assay was tested on two sets of field samples. Larval and juvenile cod were successfully detected in both tests.     

It is the economy,stupid! Projecting the fate of fish populations using ecological–economic modeling

Four marine fish species are among the most important on the world market: cod, salmon, tuna, and sea bass. While the supply of North American and European markets for two of these species – Atlantic salmon and European sea bass – mainly comes from fish farming, Atlantic cod and tunas are mainly caught from wild stocks. We address the question what will be the status of these wild stocks in the midterm future, in the year 2048, to be specific. Whereas the effects of climate change and ecological driving forces on fish stocks have already gained much attention, our prime interest is in studying the effects of changing economic drivers, as well as the impact of variable management effectiveness. Using a process‐based ecological–economic multispecies optimization model, we assess the future stock status under different scenarios of change. We simulate (i) technological progress in fishing, (ii) increasing demand for fish, and (iii) increasing supply of farmed fish, as well as the interplay of these driving forces under different scenarios of (limited) fishery management effectiveness. We find that economic change has a substantial effect on fish populations. Increasing aquaculture production can dampen the fishing pressure on wild stocks, but this effect is likely to be overwhelmed by increasing demand and technological progress, both increasing fishing pressure. The only solution to avoid collapse of the majority of stocks is institutional change to improve management effectiveness significantly above the current state. We conclude that full recognition of economic drivers of change will be needed to successfully develop an integrated ecosystem management and to sustain the wild fish stocks until 2048 and beyond.     

The fishery,biology, and management of Atlantic sturgeon,Acipenser oxyrhynchus,in North America

Synopsis The Atlantic sturgeon supported major fisheries along the entire Atlantic coast of North America. These fisheries peaked about 1890 and then suffered almost total collapse by 1905. The Atlantic sturgeon is anadromous and highly susceptible to capture during spawning migrations. Further, this species biological characteristics makes it very vulnerable to man-induced changes in natural habitat and slow to recover. Atlantic sturgeon mature at an advanced age (7–27 year for females, depending on latitude), exhibit a long interspawning period (2–5 year), and require suitable riverine, estuarine, and coastal environments for successful completion of their life cycle. Today, only remnant stocks exist in areas of former abundance. Management regulations vary considerably from state to state and range from full protection to no protection. Biological data are needed to: identify and characterize specific spawning and nursery areas; delineate migratory patterns and recruitment to various stocks; establish stock abundance; and, assess effects of various management strategies. In order to protect remaining stocks, the imposition of a total harvesting moratorium is recommended.     

Density dependence and the economic efficacy of marine reserves

Predictions on the efficacy of marine reserves for benefiting fisheries differ in large part due to considerations of models of either intra- or inter-cohort population density regulating fish recruitment. Here, I consider both processes acting on recruitment and show using a bioeconomic model how for many fisheries density dependent recruitment dynamics interact with harvest costs to influence fishery profit with reserves. Reserves consolidate fishing effort, favoring fisheries that can profitably harvest low-density stocks of species where adult density mediates recruitment. Conversely, proportion coastline in reserves that maximizes profit, and relative improvement in profit from reserves over conventional management, decline with increasing harvest costs and the relative importance of intra-cohort density dependence. Reserves never increase profit when harvest cost is high, regardless of density dependent recruitment dynamics. I quantitatively synthesize diverse results in the literature, show disproportionate effects on the economic performance of reserves from considering only inter- or intra-cohort density dependence, and highlight fish population and fishery dynamics predicted to be complementary to reserve management. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Climate,plankton and cod

While a few North Atlantic cod stocks are stable, none have increased and many have declined in recent years. Although overfishing is the main cause of most observed declines, this study shows that in some regions, climate by its influence on plankton may exert a strong control on cod stocks, complicating the management of this species that often assumes a constant carrying capacity. First, we investigate the likely drivers of changes in the cod stock in the North Sea by evaluating the potential relationships between climate, plankton and cod. We do this by deriving a Plankton Index that reflects the quality and quantity of plankton food available for larval cod. We show that this Plankton Index explains 46.24% of the total variance in cod recruitment and 68.89% of the variance in total cod biomass. Because the effects of climate act predominantly through plankton during the larval stage of cod development, our results indicate a pronounced sensitivity of cod stocks to climate at the warmer, southern edge of their distribution, for example in the North Sea. Our analyses also reveal for the first time, that at a large basin scale, the abundance of Calanus finmarchicus is associated with a high probability of cod occurrence, whereas the genus Pseudocalanus appears less important. Ecosystem‐based fisheries management (EBFM) generally considers the effect of fishing on the ecosystem and not the effect of climate‐induced changes in the ecosystem state for the living resources. These results suggest that EBFM must consider the position of a stock within its ecological niche, the direct effects of climate and the influence of climate on the trophodynamics of the ecosystem.     

Intense Habitat-Specific Fisheries-Induced Selection at the Molecular Pan I Locus Predicts Imminent Collapse of a Major Cod Fishery

Predation is a powerful agent in the ecology and evolution of predator and prey. Prey may select multiple habitats whereby different genotypes prefer different habitats. If the predator is also habitat-specific the prey may evolve different habitat occupancy. Drastic changes can occur in the relation of the predator to the evolved prey. Fisheries exert powerful predation and can be a potent evolutionary force. Fisheries-induced selection can lead to phenotypic changes that influence the collapse and recovery of the fishery. However, heritability of the phenotypic traits involved and selection intensities are low suggesting that fisheries-induced evolution occurs at moderate rates at decadal time scales. The Pantophysin I (Pan I) locus in Atlantic cod (Gadus morhua), representing an ancient balanced polymorphism predating the split of cod and its sister species, is under an unusual mix of balancing and directional selection including current selective sweeps. Here we show that Pan I alleles are highly correlated with depth with a gradient of 0.44% allele frequency change per meter. AA fish are shallow-water and BB deep-water adapted in accordance with behavioral studies using data storage tags showing habitat selection by Pan I genotype. AB fish are somewhat intermediate although closer to AA. Furthermore, using a sampling design covering space and time we detect intense habitat-specific fisheries-induced selection against the shallow-water adapted fish with an average 8% allele frequency change per year within year class. Genotypic fitness estimates (0.08, 0.27, 1.00 of AA, AB, and BB respectively) predict rapid disappearance of shallow-water adapted fish. Ecological and evolutionary time scales, therefore, are congruent. We hypothesize a potential collapse of the fishery. We find that probabilistic maturation reaction norms for Atlantic cod at Iceland show declining length and age at maturing comparable to changes that preceded the collapse of northern cod at Newfoundland, further supporting the hypothesis. We speculate that immediate establishment of large no-take reserves may help avert collapse.     

A rough guide to population change in exploited fish stocks

Interpreting how populations will change in response to exploitation is essential to the sound management of fish stocks. While deterministic models can be of use in evaluating sustainable fishing rates, the inherent variability of fish populations limits their value. In this paper a probabilistic approach is investigated which avoids having to make strong assumptions about the functional relationship between spawning stock size and the annual number of young fish (recruits) produced. Empirical probability distributions for recruits are derived, conditioned on stock size, and used to indicate likely stock changes under different fishing mortality rates. The method is applied to cod ( Gadus morhua ) in the North Sea to illustrate how population change can be inferred and used by fishery managers to choose fishing mortality rates which are likely to achieve sustainable exploitation.     

Lessons for stock assessment from the northern cod collapse

Fisheries assessment scientists can learn at least three lessons from the collapse of the northern cod off Newfoundland: (1) assessment errors can contribute to overfishing through optimistic long-term forecasts leading to the build-up of overcapacity or through optimistic assessments which lead to TACs being set higher than they should; (2) stock size overestimation is a major risk when commercial catch per effort is used as an abundance trend index, so there is continued need to invest in survey indices of abundance trend no matter what assessment methodology is used; and (3) the risk of recruitment overfishing exists and may be high even for very fecund species like cod. This implies that harvest rate targets should be lower than has often been assumed, especially when stock size assessments are uncertain. In the end, the high cost of information for accurate stock assessment may call for an alternative approach to management, involving regulation of exploitation rate via measures such as large-scale closures (refuges) that directly restrict the proportion of fish available to harvest. Development of predictive models for such regulatory options is a major challenge for fisheries assessment science.     

An Evaluation of Rebuilding Policies for U.S. Fisheries

Rebuilding depleted fish populations is a priority of modern fisheries management. In the U.S., strong statutory mandates extend to both the goals and process by which stocks are to be rebuilt. However, the National Standard Guidelines that govern the implementation of the Magnuson-Stevens Fishery Conservation and Management Act may change to increase flexibility in rebuilding requirements. In this study we evaluate performance of the status quo approach to fish stock rebuilding in the United States against 3 alternatives that have been proposed to improve rebuilding outcomes. These alternatives either simplify the analytical requirements of rebuilding analyses or apply ‘best practices’ in fisheries management, thereby avoiding the need for rebuilding analyses altogether. We use a Management Strategy Evaluation framework to evaluate rebuilding options across 6 fish life history types and 5 possible real-world fishery scenarios that include options for stock assessment quality, multiple fleets, and the degree to which the stocks are overfished at the start of the analysis. We show that the status quo rebuilding plan and a harvest control rule that reduces harvest rates at low stock size generally achieve the best rebuilding outcomes across all life-history types and fishery scenarios. Both approaches constrain fishing in the short term, but achieve high catches in the medium and long term as stocks rebuild to productive levels. These results support a growing body of literature that indicates that efforts to end overfishing early pay off in the medium- to long-term with higher cumulative catches than the alternative.     

Connectivity,persistence, and loss of high abundance areas of a recovering marine fish population in the Northwest Atlantic Ocean

In the early 1990s, the Northwest Atlantic Ocean underwent a fisheries‐driven ecosystem shift. Today, the iconic cod (Gadus morhua) remains at low levels, while Atlantic halibut (Hippoglossus hippoglossus) has been increasing since the mid‐2000s, concomitant with increasing interest from the fishing industry. Currently, our knowledge about halibut ecology is limited, and the lack of recovery in other collapsed groundfish populations has highlighted the danger of overfishing local concentrations. Here, we apply a Bayesian hierarchical spatiotemporal approach to model the spatial structure of juvenile Atlantic halibut over 36 years and three fisheries management regimes using three model parameters to characterize the resulting spatiotemporal abundance structure: persistence (similarity of spatial structure over time), connectivity (coherence of temporal pattern over space), and spatial variance (variation across the seascape). Two areas of high juvenile abundance persisted through three decades whereas two in the northeast are now diminished, despite the increased abundance and landings throughout the management units. The persistent areas overlap with full and seasonal area closures, which may act as refuges from fishing. Connectivity was estimated to be 250 km, an order of magnitude less than the distance assumed by the definition of the Canadian management units (~2,000 km). The underlying question of whether there are distinct populations within the southern stock unit cannot be answered with this model, but the smaller ~250 km scale of coherent temporal patterns suggests more complex population structure than previously thought, which should be taken into consideration by fishery management.     

Optimal harvesting of fish stocks under a time-varying discount rate

Optimal control theory has been extensively used to determine the optimal harvesting policy for renewable resources such as fish stocks. In such optimisations, it is common to maximise the discounted utility of harvesting over time, employing a constant time discount rate. However, evidence from human and animal behaviour suggests that we have evolved to employ discount rates which fall over time, often referred to as “hyperbolic discounting”. This increases the weight on benefits in the distant future, which may appear to provide greater protection of resources for future generations, but also creates challenges of time-inconsistent plans. This paper examines harvesting plans when the discount rate declines over time. With a declining discount rate, the planner reduces stock levels in the early stages (when the discount rate is high) and intends to compensate by allowing the stock level to recover later (when the discount rate will be lower). Such a plan may be feasible and optimal, provided that the planner remains committed throughout. However, in practice there is a danger that such plans will be re-optimized and adjusted in the future. It is shown that repeatedly restarting the optimization can drive the stock level down to the point where the optimal policy is to harvest the stock to extinction. In short, a key contribution of this paper is to identify the surprising severity of the consequences flowing from incorporating a rather trivial, and widely prevalent, “non-rational” aspect of human behaviour into renewable resource management models. These ideas are related to the collapse of the Peruvian anchovy fishery in the 1970's.     

Comparative study on the fatty acid and lipid composition of four marine fish larvae

• 1.1. Fatty acid and lipid class composition were determined in larvae of four marine species: Atlantic halibut (Hippoglossus hippoglossus L.), plaice (Pleuronectes platessa), cod (Gadus morhua) and turbot (Scophthalmus maximus) at hatching and prior to first feeding.
• 2.2. Total fatty acid content decreased in the four species with up to 50% reduction in one of the halibut groups. Docosahexanaoic acid (22:6 n-3) was especially utilized.
• 3.3. Low lipid utilization was found in turbot in relation to the other three species.
• 4.4. Water environmental temperature may explain some of the differences in the fatty acid utilization and the source of metabolic energy between cold water species (halibut, cod, and plaice) and temperate species (turbot), in the period from hatching to prior to first feeding.
• 5.5. Relative amounts of neutral lipids and phospholipids were similar in plaice, cod and halibut, approximately 25% and 75% of total lipids, respectively, and were approximately constant during the yolk-sac stage. Neutral lipids were dominant for turbot at hatching, accounting for 53–55% of the total lipids, while phospholipids predominated prior to first feeding, being 56–59%.
• 6.6. Phosphatidylcholine was catabolized in halibut, plaice and cod but not in turbot, while phosphatidylethanolamine tended to be synthesized in all four species.

     

Stock collapse and its effect on species interactions: Cod and herring in the Norwegian‐Barents Seas system as an example

Both the Norwegian Spring Spawning herring (Clupea harengus) and the Northeast Arctic (NEA) cod (Gadus morhua) are examples of strong stock reduction and decline of the associated fisheries due to overfishing followed by a recovery. Cod and herring are both part of the Barents Sea ecosystem, which has experienced major warming events in the early (1920–1940) and late 20th century. While the collapse or near collapse of these stocks seems to be linked to an instability created by overfishing and climate, the difference of population dynamics before and after is not fully understood. In particular, it is unclear how the changes in population dynamics before and after the collapses are associated with biotic interactions. The combination of the availability of unique long‐term time series for herring and cod makes it a well‐suited study system to investigate the effects of collapse. We examine how species interactions may differently affect the herring and cod population dynamic before and after a collapse. Particularly we explore, using a GAM modeling approach, how herring could affect cod and vice versa. We found that the effect of cod biomass on herring that was generally positive (i.e., covariation) but the effect became negative after the collapse (i.e., predation or competition). Likewise a change occurred for the cod, the juvenile herring biomass that had no effect before the collapse had a negative effect after. Our results indicate that the population collapses may alter the inter‐specific interactions and response to abiotic environmental changes. While the stocks are at similar abundance levels before and after the collapses, the system is potentially different in its functioning and may require different management action.     

Fisheries management in practice: review of 13 commercially important fish stocks

This paper comparatively reviews several commercially important fish stocks, their state and their management in various regions of the world including Japanese anchovy, Bay of Biscay anchovy, North Sea sandeel, North Sea herring, Icelandic cod, Barents Sea cod, South African cape hakes, sockeye salmon, chinook salmon, southern bluefin tuna, Pacific halibut, Greenland halibut and Patagonian toothfish. The reviewed fish stocks are systemized in three categories: (1) stock properties and status; (2) management structure and objectives; and (3) management advice. We gather evidence to outline qualities of management regimes that are recommended and highlight those that most often fail. Robust management, biological limits (reference points), implementation and consensus are critical points that separate successful and unsuccessful management regimes. We evaluate each fish stock’s management performance relative to its management objectives and current conservation issues. Furthermore, we point out the importance of stakeholder involvement in fisheries management as well as the problems that international fisheries commissions face through examples from the case studies. Management successes tended to be single-nation and single-stock fisheries with capacity control and clear stakeholder involvement. Fisheries with fleet overcapacity, unclear objectives and illegal activity characterized the case studies with management problems.     

Historical DNA reveals the demographic history of Atlantic cod (Gadus morhua) in medieval and early modern Iceland

Atlantic cod (Gadus morhua) vertebrae from archaeological sites were used to study the history of the Icelandic Atlantic cod population in the time period of 1500–1990. Specifically, we used coalescence modelling to estimate population size and fluctuations from the sequence diversity at the cytochrome b (cytb) and Pantophysin I (PanI) loci. The models are consistent with an expanding population during the warm medieval period, large historical effective population size (N_E), a marked bottleneck event at 1400–1500 and a decrease in N_E in early modern times. The model results are corroborated by the reduction of haplotype and nucleotide variation over time and pairwise population distance as a significant portion of nucleotide variation partitioned across the 1550 time mark. The mean age of the historical fished stock is high in medieval times with a truncation in age in early modern times. The population size crash coincides with a period of known cooling in the North Atlantic, and we conclude that the collapse may be related to climate or climate-induced ecosystem change.     

Effects of climate change on four New England groundfish species

Multiple groundfish stocks in New England remain depleted despite management measures that have been effective elsewhere. A growing body of research suggests that environmental change driven by increasing concentrations of carbon dioxide in the atmosphere and ocean is unfolding more rapidly in New England than elsewhere, and is an important factor in the failure of these stocks to respond to management. We reviewed research on effects of changes in temperature, salinity, dissolved oxygen, pH, and ocean currents on pelagic life stages, post-settlement life stages, and reproduction of four species in the New England groundfish fishery: Atlantic cod (Gadus morhua), haddock (Melanogrammus aeglefinus), winter flounder (Pseudopleuronectes americanus), and yellowtail flounder (Limanda ferruginea). The volume of research on cod was nearly equal to that on the other three species combined. Similarly, many more studies examined effects of temperature than other factors. The majority of studies suggest adverse outcomes, with less evidence for mixed or positive effects. However, for all of the factors other than temperature, there are more knowledge gaps than known effects. Importantly, most work to date examines impacts in isolation, but effects might combine in nonlinear ways and cause stronger reductions in stock productivity than expected. Management strategies will need to account for known effects, nonlinear interactions, and uncertainties if fisheries in New England are to adapt to environmental change.     

Biodiversity and exploitation of the main fish stocks in the Norwegian - Barents Sea ecosystem

Juvenile herring and capelin are the main stocks of plankton feeders in the Barents Sea, the cod is the dominant predator. Warm climate favours recruitment of herring and cod, but large stocks of juvenile herring hamper survival of the capelin fry. Since the early 1970s, the herring stock has been grossly overexploited, which could have led to an imbalance in the state of the predatorprey relationships in the Barents Sea. In the 1970s and early 80s, however, cod could feed on capelin which had excellent growth and recruitment conditions when the herring was absent. The consequences of the reduced herring stock were triggered in the mid 1980s, when excellent recruitment conditions for herring and cod occurred. Three abundant year classes of cod were recruited, but the herring stock was not sufficiently large to take full advantage of the favourable recruitment conditions. Given the lack of juvenile herring and a reduced capelin stock, the rapidly growing cod stock grazed down all other available prey species in the area, including its own progeny, and starved cod, seabirds and seals have in later years appeared on the north Norwegian coast. The capelin fishery collapsed, and the traditional coastal cod fisheries have been struck by the most serious crisis on record.     

Feeding habits the Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in oceanic waters of the Northern Kuril Islands and Southeast Kamchatka

This study examines the feeding habits of the Pacific cod Gadus macrocephalus in waters off the eastern coast of the northern Kuril Islands and southern Kamchatka. In November–December 1996, the cod primarily consumed fish, which made up 47.6% of the total food mass. The proportion of cephalopods, fishery offal discarded from fishing vessels, and decapods did not exceed 18.5, 17.4, and 12.2%, respectively. Among fishes, the main prey item of the cod was atka mackerel (15.4%); among cephalopods, octopus (16.8%); among fishery offal, heads of atka mackerel (14.2%); and among decapods, majid crabs (6.4%). The rather low percentage of walleye pollock (7.3%) in the cod diet was due to the decline of the east-Kamchatka walleye pollock stock.