Phenology and North Sea cod Gadus morhua L.: has climate change affected otolith annulus formation and growth?

Timing and rate of seasonal zone formation in southern North Sea cod Gadus morhua otoliths was studied. Samples were taken from two time periods, representing low and high temperature regimes. Opaque zones were laid down between January and June, in contrast with the pattern described in other published studies. Translucent zone formation started earlier in the warmer period, corresponding to peak annual sea surface temperatures, and a period of slow body growth and low metabolic activity. Translucent zone formation, however, continued once temperatures decreased and growth rate increased. It is hypothesized that translucent zone formation is triggered at a threshold of metabolic stress, and that the combined energetic requirements of reproduction, growth and migration may maintain translucent zone formation even if feeding conditions improve. Higher temperatures had a significant negative effect on the rate of translucent zone deposition, but caused a slight increase in opaque zone formation rate. The findings of this study indicate that historical otolith collections could provide key inputs into future phenological studies to improve the understanding of climate change impacts and the dynamics of otolith structure.     

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.     

Spawning stock and recruitment in North Sea cod shaped by food and climate

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.     

Lethal oxygen levels at different temperatures and the preferred temperature during hypoxia of the Atlantic cod, Gadus morhua L.

The behavioural thermoregulation of Atlantic cod Gadus morhua L. was investigated in a shuttlebox at normoxia and at three levels of hypoxia: 30, 20 and 15% oxygen saturation.
The preferred temperatures at normoxia, 30, 20 and 15% oxygen saturation were 13·9, 13·1. 10·0 and 8·8° C, respectively.
A decrease in metabolism and an increased blood oxygen affinity are among the physiological advantages of selecting a lower temperature during hypoxia. Furthermore the chances of surviving low oxygen saturations are better at low temperatures.
In natural environments, this behaviour may result in habitat shifts of fish living in heterothermal environments with changing oxygen saturations, especially in coastal areas with eutrophication, as for example the Baltic Sea.     

Temperature tracking by North Sea benthic invertebrates in response to climate change

Climate change is a major threat to biodiversity and distributions shifts are one of the most significant threats to global warming, but the extent to which these shifts keep pace with a changing climate is yet uncertain. Understanding the factors governing range shifts is crucial for conservation management to anticipate patterns of biodiversity distribution under future anthropogenic climate change. Soft‐sediment invertebrates are a key faunal group because of their role in marine biogeochemistry and as a food source for commercial fish species. However, little information exists on their response to climate change. Here, we evaluate changes in the distribution of 65 North Sea benthic invertebrate species between 1986 and 2000 by examining their geographic, bathymetric and thermal niche shifts and test whether species are tracking their thermal niche as defined by minimum, mean or maximum sea bottom (SBT) and surface (SST) temperatures. Temperatures increased in the whole North Sea with many benthic invertebrates showing north‐westerly range shifts (leading/trailing edges as well as distribution centroids) and deepening. Nevertheless, distribution shifts for most species (3.8–7.3 km yr^?1 interquantile range) lagged behind shifts in both SBT and SST (mean 8.1 km yr^?1), resulting in many species experiencing increasing temperatures. The velocity of climate change (VoCC) of mean SST accurately predicted both the direction and magnitude of distribution centroid shifts, while maximum SST did the same for contraction of the trailing edge. The VoCC of SBT was not a good predictor of range shifts. No good predictor of expansions of the leading edge was found. Our results show that invertebrates need to shift at different rates and directions to track the climate velocities of different temperature measures, and are therefore lagging behind most temperature measures. If these species cannot withstand a change in thermal habitat, this could ultimately lead to a drop in benthic biodiversity.     

Plasticity in activity and latency to explore differs between juvenile Atlantic cod Gadus morhua across a temperature gradient

In the current study activity and latency to explore, as well as the correlation of these traits, were examined in individually marked juvenile Gadus morhua at 7, 10 and 13° C. It was concluded that individual rank order of both traits was maintained across temperature but that the level of change differed between individuals.     

A century of fish growth in relation to climate change,population dynamics and exploitation

Marine ecosystems, particularly in high‐latitude regions such as the Arctic, have been significantly affected by human activities and contributions to climate change. Evaluating how fish populations responded to past changes in their environment is helpful for evaluating their future patterns, but is often hindered by the lack of long‐term biological data available. Using otolith increments of Northeast Arctic cod (Gadus morhua) as a proxy for individual growth, we developed a century‐scale biochronology (1924–2014) based on the measurements of 3,894 fish, which revealed significant variations in cod growth over the last 91 years. We combined mixed‐effect modeling and path analysis to relate these growth variations to selected climate, population and fishing‐related factors. Cod growth was negatively related to cod population size and positively related to capelin population size, one of the most important prey items. This suggests that density‐dependent effects are the main source of growth variability due to competition for resources and cannibalism. Growth was also positively correlated with warming sea temperatures but negatively correlated with the Atlantic Multidecadal Oscillation, suggesting contrasting effects of climate warming at different spatial scales. Fishing pressure had a significant but weak negative direct impact on growth. Additionally, path analysis revealed that the selected growth factors were interrelated. Capelin biomass was positively related to sea temperature and negatively influenced by herring biomass, while cod biomass was mainly driven by fishing mortality. Together, these results give a better understanding of how multiple interacting factors have shaped cod growth throughout a century, both directly and indirectly.     

Effects of freezing on length and mass measurements of Atlantic cod Gadus morhua in the Baltic Sea

An aggregated sample of 925 Atlantic cod Gadus morhua collected by four countries in different regions of the Baltic Sea during different seasons were measured (total length, L_T = 161–890 mm and weighed (mass, M = 45–6900 g) both before freezing and after defrosting. The cod were found to decrease significantly in both L_T and M following death and frozen storage. There was an average (±SD) change in L_T of −2.91% (±0.05%) following freezing, independent of starting L_T. Total M changed by −2.65% (±0.14%), independent of starting mass. Shrinkage of L_T and M did not differ significantly between 1 and 4 months frozen storage, though L_T shrinkage was significantly greater after 1 or 4 months in the freezer compared with after 5 days. There was significant variation in L_T and M shrinkage between regions of capture. A significant negative relationship between condition of cod and L_T or M change was also observed. Equations to back-calculate fresh L_T and M from thawed L_T, M and standard length (L_S), gutted L_T, gutted L_S and gutted M are provided.     

Effects of temperature, hypoxia and activity on the metabolism of juvenile Atlantic cod

Standard metabolic rate (SMR), active metabolic rate (AMR) and critical oxygen saturation ( S_crit ) were measured in Atlantic cod Gadus morhua at 5, 10 and 15° C. The SMR was 35.5, 57.0 and 78.2 mg O₂ kg⁻¹ h⁻¹ and S_crit was 16.5, 23.2 and 30.3%, at 5, 10 and 15° C, respectively. Previously reported SMR for Atlantic cod from arctic waters at 4° C was twice that measured at 5° C in the present study. A possible intraspecific latitudinal difference in the SMR is discussed. The AMR was 146.6, 197.9 and 200.4 mg O₂ kg⁻¹ h⁻¹ and the critical swimming speed ( U_crit ) was 1 6, 1.7 and 1.9 at 5, 10 and 15° C, respectively. The maximum oxygen consumption was found to be associated with exercise, rather than recovery from exercise as previously reported in another Study of Cod metabolism.     

Diet of Atlantic cod in the southern Gulf of St Lawrence as an index of ecosystem change, 1959–2000

Major temporal changes in the importance of euphausiids and Atlantic herring Clupea harengus in the diet of Atlantic cod Gadus morhua , 10–75 cm L _F, occurred in shallow waters (<100 m depths) of the southern Gulf of St Lawrence between 1959 and 2000. Euphausiids represented 6–70% of prey mass for Atlantic cod 31–60 cm L _F from 1959 to 1987 but only trace amounts were detected in stomachs collected from 1990 to 2000. Mysids and gammarid amphipods (for Atlantic cod ≤45 cm L _F), and Atlantic herring (for Atlantic cod >45 cm L _F) largely replaced euphausiids in the Atlantic cod diets from 1990 to 2000. This diet change suggested there has been a major perturbation of the food web of the southern Gulf of St Lawrence. The importance of fishes (mostly Atlantic herring) in the diet of Atlantic cod >45 cm L _F increased significantly between the periods 1959–1980 and 1987–2000. Atlantic herring comprised 0–4% (mean 1.3%) of prey mass of Atlantic cod 46–60 cm L _F from 1959 to 1980 and increased to 2–42% (mean 19.6%) of the diet from 1987 to 2000. Atlantic herring comprised 0–25% (mean 9.4%) of the prey mass of Atlantic cod 61–75 cm L _F from 1959 to 1980 and increased to 42–81% (mean 54.2%) of prey mass from 1987 to 2000. This increased consumption of Atlantic herring was consistent with observed changes in abundance of Atlantic herring in the ecosystem between the late 1970s and 2000. The large changes in consumption of euphausiids and Atlantic herring represent highly significant changes and would need to be included in the development and interpretation of ecosystem-based management models for this ecosystem.     

Mechanistic insights into the effects of climate change on larval cod

Understanding the biophysical mechanisms that shape variability in fisheries recruitment is critical for estimating the effects of climate change on fisheries. In this study, we used an Earth System Model (ESM) and a mechanistic individual‐based model (IBM) for larval fish to analyze how climate change may impact the growth and survival of larval cod in the North Atlantic. We focused our analysis on five regions that span the current geographical range of cod and are known to contain important spawning populations. Under the SRES A2 (high emissions) scenario, the ESM‐projected surface ocean temperatures are expected to increase by >1 °C for 3 of the 5 regions, and stratification is expected to increase at all sites between 1950–1999 and 2050–2099. This enhanced stratification is projected to decrease large (>5 μm ESD) phytoplankton productivity and mesozooplankton biomass at all 5 sites. Higher temperatures are projected to increase larval metabolic costs, which combined with decreased food resources will reduce larval weight, increase the probability of larvae dying from starvation and increase larval exposure to visual and invertebrate predators at most sites. If current concentrations of piscivore and invertebrate predators are maintained, larval survival is projected to decrease at all five sites by 2050–2099. In contrast to past observed responses to climate variability in which warm anomalies led to better recruitment in cold‐water stocks, our simulations indicated that reduced prey availability under climate change may cause a reduction in larval survival despite higher temperatures in these regions. In the lower prey environment projected under climate change, higher metabolic costs due to higher temperatures outweigh the advantages of higher growth potential, leading to negative effects on northern cod stocks. Our results provide an important first large‐scale assessment of the impacts of climate change on larval cod in the North Atlantic.     

Ecological forecasting under climate change: the case of Baltic cod

Good decision making for fisheries and marine ecosystems requires a capacity to anticipate the consequences of management under different scenarios of climate change. The necessary ecological forecasting calls for ecosystem-based models capable of integrating multiple drivers across trophic levels and properly including uncertainty. The methodology presented here assesses the combined impacts of climate and fishing on marine food-web dynamics and provides estimates of the confidence envelope of the forecasts. It is applied to cod (Gadus morhua) in the Baltic Sea, which is vulnerable to climate-related decline in salinity owing to both direct and indirect effects (i.e. through species interactions) on early-life survival. A stochastic food web-model driven by regional climate scenarios is used to produce quantitative forecasts of cod dynamics in the twenty-first century. The forecasts show how exploitation would have to be adjusted in order to achieve sustainable management under different climate scenarios.     

Potential effects of climate change on plant species in the Faroe Islands

Aim To identify the effect of climate change on selected plant species representative of the main vegetation types in the Faroe Islands. Due to a possible weakening of the North Atlantic Current, it is difficult to predict whether the climate in the Faroe Islands will be warmer or colder as a result of global warming. Therefore, two scenarios are proposed. The first scenario assumes an increase in summer and winter temperature of 2 °C, and the second a decrease in summer and winter temperature of 2 °C. Location Temperate, low alpine and alpine areas in the northern and central part of the Faroe Islands. Methods The responses of 12 different plant species in the Faroe Islands were tested against measured soil temperature, expressed as T_min, T_max, snow cover and growing degree days (GDD), using generalised linear modelling (GLM). Results The tolerance to changes in winter soil temperature (0.3–0.8 °C) was found to be lower than the tolerance to changing summer soil temperature (0.7–1.0 °C), and in both cases lower than the predicted climate changes. Conclusions The species most affected by a warming scenario are those that are found with a limited distribution restricted to the uppermost parts of the mountains, especially Salix herbacea, Racomitrium fasciculare, and Bistorta vivipara. For other species, the effect will mainly be a general upward migration. The most vulnerable species are those with a low tolerance, especially Calluna vulgaris, and also Empetrum nigrum, and Nardus stricta. If the climate in the Faroe Islands should become colder, the most vulnerable species are those at low altitudes. A significantly lower temperature would be expected to produce a serious reduction in the extent of Vaccinium myrtillus and Galium saxatilis. Species like Empetrum nigrum, Nardus stricta, and Calluna vulgaris may also be vulnerable. In any case, these species can be expected to migrate downwards.     

Development of 25 gene‐associated microsatellite markers of Atlantic cod (Gadus morhua L.)

Microsatellites were identified by screening 2294 GenBank entries available for Atlantic cod (Gadus morhua L.), mainly representing expressed sequence tags and cDNA sequences. Ninety‐two novel microsatellite loci (tetra‐, tri‐ and dinucleotides) were characterized on 96 individuals. This strategy yielded 25 gene‐associated polymorphic microsatellite markers (11 tri‐ and 14 dinucleotides) with two to 20 alleles and an average heterozygosity of 0.48 in the population studied (range 0.02–0.89). One marker exhibited significant homozygote excess, and one of the primer pairs amplified two linked markers. The gene identity was determined at nine of the loci, confirming the associated microsatellites as type I markers.     

Behavioral responses of Atlantic cod to sea temperature changes

Understanding responses of marine species to temperature variability is essential to predict impacts of future climate change in the oceans. Most ectotherms are expected to adjust their behavior to avoid extreme temperatures and minimize acute changes in body temperature. However, measuring such behavioral plasticity in the wild is challenging. Combining 4 years of telemetry‐derived behavioral data on juvenile and adult (30–80 cm) Atlantic cod (Gadus morhua), and in situ ocean temperature measurements, we found a significant effect of sea temperature on cod depth use and activity level in coastal Skagerrak. During summer, cod were found in deeper waters when sea surface temperature increased. Further, this effect of temperature was stronger on larger cod. Diel vertical migration, which consists in a nighttime rise to shallow feeding habitats, was stronger among smaller cod. As surface temperature increased beyond ~15°C, their vertical migration was limited to deeper waters. In addition to larger diel vertical migrations, smaller cod were more active and travelled larger distances compared to larger specimens. Cold temperatures during winter tended, however, to reduce the magnitude of diel vertical migrations, as well as the activity level and distance moved by those smaller individuals. Our findings suggest that future and ongoing rises in sea surface temperature may increasingly deprive cod in this region from shallow feeding areas during summer, which may be detrimental for local populations of the species.     

Pantophysin (Pan I) locus divergence between inshore v. offshore and northern v. southern populations of Atlantic cod in the north‐east Atlantic

More than 6000 cod Gadus morhua , sampled in coastal and offshore waters stretching from the Barents Sea down to the North Sea, were analysed for frequencies of alleles at the scnDNA pantophysin locus ( Pan I)[formerly called synaptophysin ( Syp I)]. The significant allele frequency difference between the two major stocks of cod in Norway, north‐east Arctic cod (NEAC) and Norwegian coastal cod (NCC), was upheld in all years of the investigation (1993 to 2001), and applied both to larval cod and post‐juveniles of various ages. On a north‐south axis, the appearance of a latitudinal cline of post‐juvenile (≥1 year) allele frequencies was exposed. The intermediate allele frequencies in coastal areas of northern Norway, seem to a large extent to be caused by intermingling of the two stocks, although the existence of populations of coastal cod with alternative Pan I frequencies could not be ruled out. The role of selection is yet unresolved. Depth of the sampling location seemed to have an effect on the allele frequencies and their temporal stability, while there was no indication of seasonal variation in the frequencies. Breeding structure was the most likely cause for upholding the extreme divergence in Pan I frequencies between NEAC and NCC.     

Implications of acute temperature and salinity tolerance thresholds for the persistence of intertidal invertebrate populations experiencing climate change

To predict whether populations of marine animals will persist in the face of changing climate conditions, it is informative to understand how past climate conditions have shaped present‐day tolerance thresholds. We examined 4 species of intertidal invertebrates (Nucella lamellosa, Littorina scutulata, Littorina sitkana, and Balanus glandula) inhabiting the coasts of Vancouver Island, Canada, where the east coast experiences historically warmer sea surface temperature (SST), warmer low tide (i.e., emersion) rock surface temperature (RST), and lower sea surface salinity (SSS) than the west coast. To determine if east coast populations have higher tolerance thresholds to acute stress than west coast populations, animals from 3 sites per coast were exposed to stressful temperatures and salinities in common garden experiments. Emersion temperature tolerance differed between populations only in N. lamellosa and B. glandula, tolerance thresholds being 1.4–1.5°C higher on the east coast. Water temperature tolerance differed between populations only in B. glandula and L. scutulata but was highest on the west coast. No differences in salinity tolerance were observed within any species. Thus, there is limited evidence of divergence among east and west coast populations in tolerance of acute stress despite the substantial historical differences in extreme temperature and salinity conditions between coasts. However, based on present‐day summertime SST and RST and known rates of change in these parameters, we predict present‐day tolerance thresholds would be sufficient to allow adults of these populations to tolerate extreme temperatures predicted for the next several hundred years, and that even a slow rate of change in acute tolerance thresholds might suffice to keep up with future temperature extremes.