The dynamics of Baltic fish stocks based on a multispecies stock production model

The multispecies stock‐production model of Horbowy developed in 1996 was further extended to include the unexploited part of a stock. The model was then applied to simulate stock dynamics and species interactions of cod, herring, and sprat in the Baltic from 1982 to 2001. The model indicates that there have been large declines in cod and herring biomass over the past two decades and a strong increase in sprat biomass in the 1990s. Using the extended stock‐production model, the relative changes in stock biomass were similar to the changes derived using the age‐structured multispecies model, the multispecies virtual population analysis (MSVPA). However, the production model estimates of the average predation mortality of young cod and young sprat are much lower than those derived from MSVPA, although the estimates for young and adult herring and adult sprat are similar in both approaches. The estimates of food suitability show that the preferred food of adult cod is adult sprat and young herring, while the suitability of young sprat, young cod, and adult herring is much smaller. The simulations performed show that the multispecies production model, which is less data‐demanding than age‐structured MSVPA, can provide estimates of stock dynamics and species interactions that are largely consistent with those estimated by MSVPA. The quality of input data in terms of recruitment and fishing‐effort indices strongly impacts the reliability of the model's results.     

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.     

Temporal Dynamics of Top Predators Interactions in the Barents Sea

The Barents Sea system is often depicted as a simple food web in terms of number of dominant feeding links. The most conspicuous feeding link is between the Northeast Arctic cod Gadus morhua, the world''s largest cod stock which is presently at a historical high level, and capelin Mallotus villosus. The system also holds diverse seabird and marine mammal communities. Previous diet studies may suggest that these top predators (cod, bird and sea mammals) compete for food particularly with respect to pelagic fish such as capelin and juvenile herring (Clupea harengus), and krill. In this paper we explored the diet of some Barents Sea top predators (cod, Black-legged kittiwake Rissa tridactyla, Common guillemot Uria aalge, and Minke whale Balaenoptera acutorostrata). We developed a GAM modelling approach to analyse the temporal variation diet composition within and between predators, to explore intra- and inter-specific interactions. The GAM models demonstrated that the seabird diet is temperature dependent while the diet of Minke whale and cod is prey dependent; Minke whale and cod diets depend on the abundance of herring and capelin, respectively. There was significant diet overlap between cod and Minke whale, and between kittiwake and guillemot. In general, the diet overlap between predators increased with changes in herring and krill abundances. The diet overlap models developed in this study may help to identify inter-specific interactions and their dynamics that potentially affect the stocks targeted by fisheries.     

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.     

Shifting dynamic forces in fish stock fluctuations triggered by age truncation?

Accumulating evidence shows that environmental fluctuations and exploitation jointly affect marine fish populations, and understanding their interaction is a key issue for fisheries ecology. In particular, it has been proposed that age truncation induced by fisheries exploitation may increase the population's sensitivity to climate. In this study, we use unique long‐term abundance data for the Northeast Arctic stock of cod (Gadus morhua) and the Norwegian Spring‐Spawning stock of herring (Clupea harengus), which we analyze using techniques based on age‐structured population matrices. After identifying time periods with different age distributions in the spawning stock, we use linear models to quantify the relative effect of exploitation and temperature on the population growth rates. For the two populations, age truncation was found to be associated with an increasing importance of temperature and a relatively decreasing importance of exploitation, while the population growth rate became increasingly sensitive to recruitment variations. The results suggested that the removal of older age classes reduced the buffering capacity of the population, thereby making the population growth rate more dependent on recruitment than adult survival and increasing the effect of environmental fluctuations. Age structure appeared as a key characteristic that can affect the response of fish stocks to climate variations and its consequences may be of key importance for conservation and management.     

Timing and abundance as key mechanisms affecting trophic interactions in variable environments

Climatic changes are disrupting otherwise tight trophic interactions between predator and prey. Most of the earlier studies have primarily focused on the temporal dimension of the relationship in the framework of the match–mismatch hypothesis. This hypothesis predicts that predator's recruitment will be high if the peak of the prey availability temporally matches the most energy‐demanding period of the predators breeding phenology. However, the match–mismatch hypothesis ignores the level of food abundance while this can compensate small mismatches. Using a novel time‐series model explicitly quantifying both the timing and the abundance component for trophic relationships, we here show that timing and abundance of food affect recruitment differently in a marine (cod/zooplankton), a marine–terrestrial (puffin/herring) and a terrestrial (sheep/vegetation) ecosystem. The quantification of the combined effect of abundance and timing of prey on predator dynamics enables us to come closer to the mechanisms by which environment variability may affect ecological systems.     

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.     

Fish, Fact and Fantasy: a Long View

This paper suggests ways forward from the widely perceived present failures of fishery assessment and management. A history of fishery yield modelling is presented from the carefree days of the 1950s to the depressing series of stock collapses and depletions of the 1980s. Underlying this gruesome story has been the failure of management by quotas to arrest overcapacity in fishing power, the lack of robust and informative reference points and the inadequacy of methods dealing with some multispecies fisheries. The paper refines the use of the concept of Fext, defined as the minimum value of F in a self-regenerating yield model that leads to eventual extinction in a family of yield curves generated with a range of stock recruitment curves. Model reconstructions for North Sea cod and Icelandic herring make evident calamitous losses in catches forgone as result of the failure of rational management. An optimistic agenda that may achieve more effective fishery management in the future is presented. In some ways, we may have been trying to be too clever. A simple management system based on careful monitoring of fishing effort, biological targets such as F95, and exploitation of a diversity of fish resources may suffice to avert further disaster and hedge against uncertainty.     

Recruitment prediction for Pacific herring (Clupea pallasi) on the west coast of Vancouver Island, Canada

The accurate prediction of recruitment to the fishery is a very important tool within the management structure of any fish stock being exploited. In the case of the Pacific herring, Clupea pallasi, fishery in Canada, a forecast of the abundance of each herring stock is particularly important for formulating an annual catch quota. The sustainable management of the fishery and the resource is based in part on accurate recruitment forecasting because Pacific herring are short-lived and so the recruitment contributes a significant part of the total spawning run targeted by the fishery each year. Several factors are believed be important in determining the success of recruitment besides spawners biomass. Since herrings are “r” strategists, conditions related to the egg, the planktonic, or even the juvenile stage might determine the future level of recruitment. Recently a formula that defines conditions for a semi-quantitative level of recruitment forecast was elaborated using genetic algorithms and current study attempts to improve on this model. Using salinity in two quarterly periods during the planktonic and pre-recruit stages, temperature and spawning biomass for the west coast of Vancouver Island stock, classification rules that define recruitment in 3 different levels (low, medium and high) were developed with a genetic algorithm, setting low and high boundaries for each condition. A 75% success in classifying recruitment was obtained. The model was shown to be particularly effective at predicting when the recruitment would be low, which could be important from the perspective of the Precautionary Approach and the sustainable management of this stock.     

An analysis of a nonlinear stage-structured cannibalism model with application to the Northeast Arctic cod stock

A two-dimensional stage-structured population model with nonlinear cannibalism terms is studied. We show that there is a large parameter interval where the nontrivial equilibrium of the model is the only stable attractor, but that there also exist parameter intervals where we find quasiperiodic, periodic and chaotic dynamics. Moreover, in the interplay between increasing the fecundity and increasing the cannibalism pressure, the former turns out to be a destabilizing effect while the latter tends to act in a stabilizing fashion. Finally, we have applied the model to the North Atlantic cod stock using ICES biomass estimates. Our main conclusion from this study is that the combined effect of recruitment and cannibalism may not serve as an explanation of the observed fluctuations in the cod stock.     

Extrapolating predation mortalities back in time: an example from North-east Arctic cod cannibalism

Cannibalism is known to be a significant source of natural mortality of young North-east Arctic (NEA) cod. Cannibalism data, starting from 1984, have been used in NEA cod stock assessments since 1995, which has led to inconsistency in the cod abundance time series from 1946 to the present. To address this inconsistency, this study estimates the cannibalism-induced mortality (M2) of NEA cod at age 3–5 for the period 1946–1983. Combined qualitative and quantitative cod stomach content data for 1984–2010 were used to make the M2 estimations for age groups 3–5 (ICES 2014), then different factors including SSB were used to examine which covariates explained variability in M2 and thus make predictions for 1946–1983. The level of cannibalism was estimated to be high in the 1950s – early1960s. VPA-based assessment was run using these estimated M2 values. As a result, numbers of cod eaten by their conspecifics in the historical period and new increased recruitment estimates at age 3 were computed. The main factors affecting cannibalism appeared to be young cod abundance, total stock biomass (TSB) of large cod, and capelin total stock biomass (which represents an alternative prey). The problems involved in using the new recruitment time series in fishery management are discussed. The methodology presented here represents a generic approach to extending predation mortalities back in time to improve historical stock estimates.     

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.     

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.     

Characteristics of the Norwegian Coastal Current during Years with High Recruitment of Norwegian Spring Spawning Herring (Clupea harengus L.)

Norwegian Spring Spawning herring (NSSH) Clupea harengus L. spawn on coastal banks along the west coast of Norway. The larvae are generally transported northward in the Norwegian Coastal Current (NCC) with many individuals utilizing nursery grounds in the Barents Sea. The recruitment to this stock is highly variable with a few years having exceptionally good recruitment. The principal causes of recruitment variability of this herring population have been elusive. Here we undertake an event analysis using data between 1948 and 2010 to gain insight into the physical conditions in the NCC that coincide with years of high recruitment. In contrast to a typical year when northerly upwelling winds are prominent during spring, the years with high recruitment coincide with predominantly southwesterly winds and weak upwelling in spring and summer, which lead to an enhanced northward coastal current during the larval drift period. Also in most peak recruitment years, low-salinity anomalies are observed to propagate northward during the spring and summer. It is suggested that consistent southwesterly (downwelling) winds and propagating low-salinity anomalies, both leading to an enhanced northward transport of larvae, are important factors for elevated recruitment. At the same time, these conditions stabilize the coastal waters, possibly leading to enhanced production and improved feeding potential along the drift route to Barents Sea. Further studies on the drivers of early life history mortality can now be undertaken with a better understanding of the physical conditions that prevail during years when elevated recruitment occurs in this herring stock.     

Forecasting future recruitment success for Atlantic cod in the warming and acidifying Barents Sea

Productivity of marine fish stocks is known to be affected by environmental and ecological drivers, and global climate change is anticipated to alter recruitment success of many stocks. While the direct effects of environmental drivers on fish early life stage survival can be quantified experimentally, indirect effects in marine ecosystems and the role of adaptation are still highly uncertain. We developed an integrative model for the effects of ocean warming and acidification on the early life stages of Atlantic cod in the Barents Sea, termed SCREI (Simulator of Cod Recruitment under Environmental Influences). Experimental results on temperature and CO₂ effects on egg fertilization, egg and larval survival and development times are incorporated. Calibration using empirical time series of egg production, temperature, food and predator abundance reproduces age‐0 recruitment over three decades. We project trajectories of recruitment success under different scenarios and quantify confidence limits based on variation in experiments. A publicly accessible web version of the SCREI model can be run under www.oceanchange.uni-bremen.de/ ;SCREI. Severe reductions in average age‐0 recruitment success of Barents Sea cod are projected under uncompensated warming and acidification toward the middle to end of this century. Although high population stochasticity was found, considerable rates of evolutionary adaptation to acidification and shifts in organismal thermal windows would be needed to buffer impacts on recruitment. While increases in food availability may mitigate short‐term impacts, an increase in egg production achieved by stock management could provide more long‐term safety for cod recruitment success. The SCREI model provides a novel integration of multiple driver effects in different life stages and enables an estimation of uncertainty associated with interindividual and ecological variation. The model thus helps to advance toward an improved empirical foundation for quantifying climate change impacts on marine fish recruitment, relevant for ecosystem‐based assessments of marine systems under climate change.     

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.     

Local and large-scale climatic variables as predictors of the breeding numbers of endangered Lesser Black-backed Gulls on the Norwegian Coast

In the 1970s and 1980s, the nominate subspecies of the Lesser Black-backed Gull (Larus fuscus fuscus) showed a dramatic drop in breeding numbers on the Norwegian Coast, and in 2000, the population in some colonies was only 10–20% of the population in 1980. This decline has been attributed to the collapse in the stock of Norwegian spring spawning herring (Clupea harengus). In this study, we examined whether local climate (sea and air temperatures), winter NAO (North Atlantic Oscilliation), and the year-class strength and size of 0-group herring could predict the relative changes in breeding numbers between years, mainly after the population collapse. Breeding birds were counted in 19 of the years between 1980 and 2007 in an archipelago on the coast of Helgeland, northern Norway. The best model predicting changes in breeding numbers for the period between 1980 and 2005 (for which data on 0-group herring was available) included mean local air temperature in winter (January–March) and winter NAO, explaining 57% of the variation between years, while the other factors had little effect. When also adding the years 2006–2007 (no herring data), the best model included only mean air temperature in winter, explaining 41% of the variation. In conclusion, the high positive correlation between breeding numbers and climatic factors probably resulted from a higher availability of important fish prey after mild winters, for which 0-group herring presently may only account for a limited proportion. However, this prey might have been of much more importance prior to the population decline.     

Ecological significance of 0-group fish in the Barents Sea ecosystem

Investigations into the 0-group fish in the Barents Sea have been carried out since 1965, with the goal of estimating the abundance of 0-group fish. 0-group abundance indices have been used in the assessment of the recruitment level and in recruitment variability studies. However, the ecological importance of the 0-group fish in the Barents Sea has been less studied. Although 0-group capelin, herring, cod and haddock are widely distributed in the Barents Sea, the central area seems to be the most important, accounting for approximately 50–80% of the annual biomass. The total biomass of the four most abundant 0-group fish species can be up to 3.3 million tonnes, with an average of 1.3 million tonnes (1993–2009). Wide distribution and high biomass of pelagically distributed 0-group fish make these fishes an important element in the energy transport between different trophic levels and different geographical areas, having a critical impact on the entire Barents Sea ecosystem. In recent years, capelin have shown a pronounced northward shift in biomass distribution, and several successive strong year classes occurred during warm temperature conditions. Cod biomasses were unexpectedly low during warm years and were positively correlated with spawning stock biomass, while the correlation with temperature was not significant. Haddock and herring show, as expected, increasing biomass with increased temperature when the spawning stock is at a sufficiently high level.     

The origins of intensive marine fishing in medieval Europe: the English evidence

The catastrophic impact of fishing pressure on species such as cod and herring is well documented. However, the antiquity of their intensive exploitation has not been established. Systematic catch statistics are only available for ca.100 years, but large-scale fishing industries existed in medieval Europe and the expansion of cod fishing from the fourteenth century (first in Iceland, then in Newfoundland) played an important role in the European colonization of the Northwest Atlantic. History has demonstrated the scale of these late medieval and post-medieval fisheries, but only archaeology can illuminate earlier practices. Zooarchaeological evidence shows that the clearest changes in marine fishing in England between AD 600 and 1600 occurred rapidly around AD 1000 and involved large increases in catches of herring and cod. Surprisingly, this revolution predated the documented post-medieval expansion of England's sea fisheries and coincided with the Medieval Warm Period--when natural herring and cod productivity was probably low in the North Sea. This counterintuitive discovery can be explained by the concurrent rise of urbanism and human impacts on freshwater ecosystems. The search for 'pristine' baselines regarding marine ecosystems will thus need to employ medieval palaeoecological proxies in addition to recent fisheries data and early modern historical records.