Climate change and fishing: a century of shifting distribution in North Sea cod

Globally, spatial distributions of fish stocks are shifting but although the role of climate change in range shifts is increasingly appreciated, little remains known of the likely additional impact that high levels of fishing pressure might have on distribution. For North Sea cod, we show for the first time and in great spatial detail how the stock has shifted its distribution over the past 100 years. We digitized extensive historical fisheries data from paper charts in UK government archives and combined these with contemporary data to a time‐series spanning 1913–2012 (excluding both World Wars). New analysis of old data revealed that the current distribution pattern of cod – mostly in the deeper, northern‐ and north‐easternmost parts of the North Sea – is almost opposite to that during most of the Twentieth Century – mainly concentrated in the west, off England and Scotland. Statistical analysis revealed that the deepening, northward shift is likely attributable to warming; however, the eastward shift is best explained by fishing pressure, suggestive of significant depletion of the stock from its previous stronghold, off the coasts of England and Scotland. These spatial patterns were confirmed for the most recent 3½ decades by data from fisheries‐independent surveys, which go back to the 1970s. Our results demonstrate the fundamental importance of both climate change and fishing pressure for our understanding of changing distributions of commercially exploited fish.     

Synergistic Effects of Climate and Fishing in a Marine Ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere. Author Contributions  RRK conceived the project and GB analysed the data. RRK, GB and JAL co-wrote the paper.     

Environmental influences on long-term variability in marine plankton

The major patterns of geographical and seasonal variability of the plankton of the north-east Atlantic and the North Sea are described to provide the background to a presentation of the dominant patterns of year-to-year fluctuations in the abundance of the plankton of the area for the period 1948 to 1984. A feature of the variability is a marked similarity both between species and between areas. The main pattern of year-to-year change has the form of a quasi linear downward trend in abundance with, superimposed on this, an element of variability with a periodicity of about three years. There is a complex relationship between the plankton and an estimate of changes in the frequency of westerly weather which can be interpreted in terms of influences acting over limited periods of the seasonal cycle coupled with persistence in the stocks of zooplankton. Relationships between year-to-year variations in the abundance of phytoplankton and zooplankton can be interpreted in terms of a response by the zooplankton to variations in food supply coupled with feed-back to the phytoplankton involving in situ nutrient regeneration.     

North Sea cod and climate change – modelling the effects of temperature on population dynamics

In order to examine the likely impacts of climate change on fish stocks, it is necessary to couple the output from large‐scale climate models to fisheries population simulations. Using projections of future North Sea surface temperatures for the period 2000–2050 from the Hadley General Circulation Model, we estimate the likely effects of climate change on the North Sea cod population. Output from the model suggests that increasing temperatures will lead to an increased rate of decline in the North Sea cod population compared with simulations that ignore environmental change. Although the simulation developed here is relatively simplistic, we demonstrate that inclusion of environmental factors in population models can markedly alter one's perception of how the population will behave. The development of simulations incorporating environment effects will become increasingly important as the impacts of climate change on the marine ecosystem become more pronounced.     

Evidence for selection and spatially distinct patterns found in a putative zona pellucida gene in Pacific cod,and implications for management

Genetic differentiation has been observed in marine species even when no obvious barriers to gene flow exist, and understanding such differentiation is essential for effective fisheries management. Highly differentiated outlier loci can provide information on how genetic variation might not only contribute to local adaptation but may also be affected by historical demographic events. A locus which aligned to a predicted zona pellucida sperm‐binding protein 3 gene (ZP3) in Atlantic cod (Gadus morhua) was previously identified as the highest outlier based on F _ST in a RADseq study of Pacific cod (Gadus macrocephalus) across the West Coast of North America. However, because of the limited length of the RAD sequence and restricted geographic area of sampling, no conclusion on the functional significance of the observed variation was possible. In other marine species, ZP3 is involved in reproductive isolation, local adaptation, and has neofunctionalized as an antifreeze gene, and so it may provide important insights in functional population structure of Pacific cod. Here, we sequenced a 544‐bp region of ZP3 in 230 Pacific cod collected from throughout their geographic range. We observed striking patterns of spatial structuring of ZP3 haplotypes, with a sharp break near Kodiak, Alaska, USA where populations within ~200 km of each other are nearly fixed for different haplotypes, contrasting a pattern of isolation by distance at other genetic markers in this region (F _ST = 0.003). Phylogenetic analysis of ZP3 haplotypes revealed that the more southern haplotypes appear to be ancestral, with the northern haplotype evolving more recently, potentially in response to a novel selective pressure as Pacific cod recolonized northern latitudes after glaciation. The sharp break in haplotype frequencies suggests strong selective pressures are operating on small spatial scales and illustrates that selection can create high divergence even in marine species with ample opportunities for gene flow.     

Gene flow and lack of population differentiation in Atlantic cod, Gadus morhua L., from Iceland, and comparison of cod from Norway and Newfoundland

Restriction analysis of mitochondrial DNA was used to study genetic variation and geographic population structure of Atlantic cod from localities around Iceland. Gene phylogenies were constructed and geographic locations superimposed on these. The variation was not localized. Estimated gene flow was large. Thus, Atlantic cod in Iceland belong to a single genetic population. Analyses of published sequence variation of cod from Norway and Newfoundland showed that the extensive continuity of intraspecific phylogeny of cod in Iceland, which is very similar to other marine organisms with a similar life history, extends from Norway to Newfoundland and possibly to larger areas of the Atlantic.     

The dynamics of exploited marine fish populations and Humpty Dumpty: similarities and differences

Marine fish populations of the global oceans and particularly large-bodied, continental shelf-dwelling groundfish species of the North Atlantic, such as cod (Gadus morhua), have been strongly perturbed by over-fishing, frequently beyond levels that may have altered their capacity to recover. Age and size structure, spatial structure, reproductive potential, and other traits that convey fitness advantages are commonly lost when prolonged and excessive fishing pressure is exerted. Fisheries management protocols implemented to recover collapsed populations have been numerous and varied with all attempting to reduce or eliminate fishing pressure. Such measures, employed singly or in multiple combinations, typically involve quota reductions or fishing moratoria, area closures and other technical measures, as well as enhanced enforcement of fishing practices. A striking geographic pattern exists in the efficacy of such measures to regain lost population attributes and hence recovery. Some regional populations have recovered while others, despite management interventions lasting decades, notably, but not exclusively, those aimed at cod populations of the Northwest Atlantic, have yet to fully recover, an endpoint analogous to the conclusion of the famous nursery rhyme of Humpty Dumpty. Here we examine the dynamics of multiple collapsed populations exhibiting varying responses to recovery initiatives from the perspective of the Humpty Dumpty metaphor.     

Effect of turbidity on habitat preference of juvenile Atlantic cod, Gadus morhua

We examined the effects of turbidity on habitat preference of juvenile Atlantic cod in the laboratory, using a shuttle box where fish could select between two different habitats. In the first experiment, we compared three turbidity levels of kaolin (3, 8 and 21 beam attenuation m⁻¹). In the second experiment, we looked at the effect of turbidity media (kaolin versus algae), after controlling for spectral differences between turbidity media. Although cod preferred an intermediate turbidity of kaolin over low turbidity water, comparisons between low and high turbidity, and intermediate and high turbidity did not significantly influence habitat preference. Algae did not influence habitat preference by cod. Although other studies have found that turbidity affects both foraging and antipredator behaviour of juvenile cod, this study has shown that gradients in turbidity per se do not have a strong effect on their habitat preference.     

Causes and projections of abrupt climate-driven ecosystem shifts in the North Atlantic

Warming of the global climate is now unequivocal and its impact on Earth' functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.     

A moorable,automated plankton sampler

A moorable, automated plankton sampler is described, designed to obtain series of plankton samples over extended time periods in the absence of a surface vessel. The sampler consists of one or more net-containing boxes, a unit for generating as well as measuring water flow through the net boxes, a programmeable control unit, and a frame. Each net box contains several nets on a vinyl strip, used to move the nets from the storage chamber into the fishing position and then into the preservative chamber. Preservation of samples is in a formalin-brine mixture.Net boxes can be added and the number of nets per box changed; the prototype is described with 1 net box with 10 nets. Volume of water to be filtered, time between sample collections, number of net boxes and the number of nets in each box is programmed into the control unit before sampler deployment. Collections made with the sampler are compared with those made with a SCOR net and a pump.     

The feeding, growth and behaviour of juvenile cod, Gadus morhua L., in cold environments

Most studies on feeding-related activities of fish in cold water (0° C) have not used eurythermal species or examined a broad suite of activities. In this study we report on the feeding, growth, behaviour and conversion-efficiency of 0 + Atlantic cod in response to water temperature and food availability. In one experiment, activity, opercular beat and growth decreased with decreasing temperature over the range 8·3–0·6° C. However, the conversion efficiency increased with decreasing temperature, a result partly explained by the relative change in activity and opercular rate. In another experiment, temperature had a linear effect on growth in cod fed different food rations. Both food availability and temperature were found to influence growth in group 0 + cod. The specific growth rate of cod in this experiment exceeded 2% day^-1. Growth data collected from cod sampled in the field during the cold-water period showed a progressive increase of mean length over the season. Overall, these results demonstrate that 0 + cod have adapted to life in cold-water environments, in contrast to adult cod which mostly inhabit warmer environments.     

Modelling regional differences in 'sealworm', Pseudoterranova decipiens (Nematoda, Ascaridoidea), infections in some North Atlantic cod, Gadus morhua, stocks

The ascaridoid nematode Pseudoterranova decipiens has a complex life-cycle. Adult worms reproduce in the stomach of grey seals, Halichoerus grypus , and other pinnipeds, and the larvae are found encysted in the musculature of Atlantic cod, a common host. An increase in infection levels in Canadian, Icelandic and Norwegian cod stocks over the last decade has been attributed to recent increases in seal population numbers. A simple model is proposed for the accumulation of larval parasites in the fish through its food. The model describes how regional differences in fish growth can also explain some differences in infection levels.     

Movers and shakers: nutrient subsidies and benthic disturbance predict biofilm biomass and stable isotope signatures in coastal streams

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A trans‐Atlantic examination of haddock Melanogrammus aeglefinus food habits

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Fishing,predation, and temperature drive herring decline in a large marine ecosystem

Since 1960, landings of Atlantic herring have been the greatest of any marine species in Canada, surpassing Atlantic cod and accounting for 24% of the total seafood harvested in Atlantic Canada. The Scotian Shelf‐Bay of Fundy herring fisheries (NAFO Division 4VWX) is among Canada''s oldest and drives this productivity, accounting for up to 75% of the total herring catch in some years. The stocks’ productivity and overall health have declined since 1965. Despite management measures to promote recovery implemented since 2003, biomass remains low and is declining. The factors that drive the productivity of 4VWX herring are primarily unresolved, likely impeding the effectiveness of management actions on this stock. We evaluated potential drivers of herring variability by analyzing 52 time‐series that describe the temporal and spatial evolution of the 4VWX herring population and the physical, ecological, and anthropogenic factors that could affect them using structural equation models. Variation in herring biomass was best accounted for by the exploitation rate''s negative effect and the geographic distribution of fishing and recruitment. Thermal phenology and temperature adversely and egg predation positively impacted the early life stage mortality rate and, ultimately, adult biomass. These findings are broadly relevant to fisheries management, but particularly for 4VWX herring, where the current management approach does not consider their early life stage dynamics or assess them within the ecosystem or climate change contexts.     

Comparison of various potential fecundity models for north-east Arctic cod Gadus morhua, L. using oocyte diameter as a standardizing factor

To build a better data foundation for recruitment models of north-east Arctic cod Gadus morhua the construction of fecundity models reflecting variation in the nutritional status of the fish was attempted. The models were based on fecundity time series covering 9 years within the period 1986–2004 and included both general and year-specific approaches. Initial data analysis revealed that the potential fecundity ( F _P) (standing stock of vitellogenic oocytes) was significantly reduced as the vitellogenic oocytes increased in size towards the start of spawning. Histological examination strongly indicated that this seasonal reduction was caused by atresia. Regression analysis showed that the F _P was positively correlated to fish total length ( L _T) and the Fulton's condition factor ( K ). A multiple regression including data for all years using fish L _T, K and mean oocyte diameter ( D _O) as independent predictors described the F _P with an r ²= 0·94. This was considerably higher than comparable univariate L _T or mass-based regressions. These univariate regressions had fairly high r ² values when split by years, but not as high as found for year-specific multiple regressions. An important application for individual-based fecundity models may be to generate outputs that can be fed into stock level fecundity and recruitment models. Overall, the multivariate models seemed to be the most accurate. The multivariate model including mean D _O, however, also had the potential to correct for maturity and thus provide unbiased fecundity comparisons between years, stocks and locations.     

Characterization of 18 new microsatellite loci in Atlantic cod (Gadus morhua L.)

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