Fluctuation in stock properties of north‐east Arctic cod related to long‐term environmental changes

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Abstract From a historic perspective, the north‐east Arctic cod stock, which is found in the Barents Sea–Svalbard region, has been the most productive gadoid stock in the Atlantic. Variation in catch has always been large, but during the last 10–15 years catch and stock abundance have reached the lowest level on record. Three major causes of variation have been discussed: (i) stock reduction through exploitation; (ii) environmental influences on recruitment; and (iii) species interaction effects on maturation, growth and mortality. In addition, interactions among these three sources might be important. The influence of each specific factor is difficult to evaluate from incidental observations and short‐term time series. In this respect, the time series on catches and on biological and environmental information of this stock, which partly extend back to the 19th century, occupy a unique position in comparison to data on most other stocks. In this paper, fluctuations in catch and stock abundance are compared with changes in recruitment, size/age and growth. This information is discussed in view of historic variation in ecology and environment. The stock has been under particularly high exploitation pressure since the mid‐1970s. Further, large changes in growth rates and poor recruitment to the commercially exploited stock are characteristic of late 1980s and throughout the 1990s. The analysis shows that substantial long‐term variation might underlie short‐term variability, and more importantly, that long‐term changes roughly coincide with similar fluctuations in the environment. Such factors might substantially affect the relationship between spawning stock and recruitment, which is also apparent from the difference in conclusions reached by various published studies. Consequently, it is suggested that using a steady‐state perspective for the population dynamics may lead to mismanagement and to a reduction of the long‐term yield from this stock.

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CitationsShowing 10 of 52 papers
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Accelerated extractions of North Atlantic cod and herring, 1520–1790
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Abstract We propose the concept of Accelerated Marine Extraction to signify two periods when rapidly increasing cod (Gadus morhua, Gadidae) and herring (Clupea harengus, Clupeidae) fisheries, c.1540–1600 and c. 1730–1790, exceeded human demographic growth. Total landings vastly exceeded previous assessments and more than doubled between 1520 and 1620 from about 220,000 metric tonnes (t) to 460,000 t. Supplies of cod and herring to the European market peaked in 1788 at more than 1 million t before the unrest connected with the French Revolution brought many fisheries to a temporary halt. Accelerated Marine Extractions increased European food security at times of human demographic growth by almost doubling the supplies of fish protein per capita. While herring was the most important species by 1520, cod dominated through the period 1540–1790, and the trajectories of cod and herring extractions differed significantly. Cod landings increased almost ten‐fold between 1520 and 1790, driven by strong and sustained landings in the Northwest Atlantic. Herring landings remained stable through the 16th century but declined severely through the next 150 years. However, from 1750, herring landings quadrupled, largely because of Swedish west coast fisheries. The results fundamentally shift our understanding of the scale of Atlantic fisheries in the past and underline the role of marine resources for European societies.

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Food web dynamics affect Northeast Arctic cod recruitment.
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Proper management of ecosystems requires an understanding of both the species interactions as well as the effect of climate variation. However, a common problem is that the available time-series are of different lengths. Here, we present a general approach for studying the dynamic structure of such interactions. Specifically, we analyse the recruitment of the world's largest cod stock, the Northeast Arctic cod. Studies based on data starting in the 1970-1980s indicate that this stock is affected by temperature through a variety of pathways. However, the value of such studies is somewhat limited by the fact that they are based on a quite specific ecological and climatic situation. Recently, this stock has consisted of fairly young fish and the spawning stock has consisted of relatively few age groups. In this study, we develop a model for the effect of capelin (the cod's main prey) and herring on cod recruitment since 1973. Based on this model, we analyse data on cod, herring and temperature going back to 1921 and find that food-web effects explain a significant part of the cod recruitment variation back to around 1950.

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Does increasing mortality change the response of fish populations to environmental fluctuations?
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  • Ecology Letters
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Fluctuations of fish populations abundances are shaped by the interplay between population dynamics and the stochastic forcing of the environment. Age-structured populations behave as a filter of the environment. This filter is characterised by the species-specific life cycle and life-history traits. An increased mortality of mature individuals alters these characteristics and may therefore induce changes in the variability of populations. The response of a generic age-structured model was analysed to investigate the expected changes in the fluctuations of fish populations in response to decreased adult survival. These expectations were then tested on an extensive dataset. In accordance with theory, the analyses revealed that decreased adult survival and mean age of spawners were linked to an increase in the relative importance of short-term fluctuations. It suggests that intensive exploitation can lead to a change in the variability of fish populations, an issue of central interest from both conservation and management perspectives.

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Pronounced long-term juvenation in the spawning stock of Arcto-Norwegian cod (Gadus morhua) and possible consequences for recruitment
  • Mar 1, 2008
  • Canadian Journal of Fisheries and Aquatic Sciences
  • Geir Ottersen

The oldest and largest individuals are disappearing from many fish stocks worldwide as a result of overexploitation. This has been suggested to impair recruitment through decreasing the reproductive capacity of the spawners and increasing the mortality rate of the offspring. By using a time series on spawners biomass by age class for Arcto-Norwegian cod (Gadus morhua) from 1913–2004, I have documented pronounced changes in the spawning stock, including a trend towards younger fish, a less diverse distribution across ages, and a declining proportion of repeat spawners. Despite the total spawning stock biomass (SSB) being at similar levels now as in 1933, the mean age in the SSB has declined from 10–12.5 years to 7–8 years during the study period, and the percentage of fish of age 10 or above in the SSB has decreased from ~97% to ~10%. Contrary to earlier theoretical and experimental studies, no clear link between age structure and recruitment was found here. Recruitment to the Arcto-Norwegian cod stock may thus be more robust towards spawner juvenation than expected, possibly because of strong recruitment compensation.

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Detecting regime shifts in fish stock dynamics
  • Nov 1, 2015
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  • Tommi Perälä + 1 more

Environmental factors such as water temperature, salinity, and the abundance of zooplankton can have major effects on certain fish stocks’ ability to produce juveniles and, thus, stock renewal ability. This variability in stock productivity manifests itself as different productivity regimes. Here, we detect productivity regime shifts by analyzing recruit-per-spawner time series with Bayesian online change point detection algorithm. The algorithm infers the time since the last regime shift (change in mean or variance or both) as well as the parameters of the data-generating process for the current regime sequentially. We demonstrate the algorithm’s performance using simulated recruitment data from an individual-based model and further apply the algorithm to stock assessment estimates for four Atlantic cod (Gadus morhua) stocks obtained from RAM legacy database. Our analysis shows that the algorithm performs well when the variability between the regimes is high enough compared with the variability within the regimes. The algorithm found several productivity regimes for all four cod stocks, and the findings suggest that the stocks are currently in low productivity regimes, which have started during the 1990s and 2000s.

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Roles of density-dependent growth and life history evolution in accounting for fisheries-induced trait changes
  • Dec 9, 2016
  • Proceedings of the National Academy of Sciences
  • Anne Maria Eikeset + 5 more

The relative roles of density dependence and life history evolution in contributing to rapid fisheries-induced trait changes remain debated. In the 1930s, northeast Arctic cod (Gadus morhua), currently the world's largest cod stock, experienced a shift from a traditional spawning-ground fishery to an industrial trawl fishery with elevated exploitation in the stock's feeding grounds. Since then, age and length at maturation have declined dramatically, a trend paralleled in other exploited stocks worldwide. These trends can be explained by demographic truncation of the population's age structure, phenotypic plasticity in maturation arising through density-dependent growth, fisheries-induced evolution favoring faster-growing or earlier-maturing fish, or a combination of these processes. Here, we use a multitrait eco-evolutionary model to assess the capacity of these processes to reproduce 74 y of historical data on age and length at maturation in northeast Arctic cod, while mimicking the stock's historical harvesting regime. Our results show that model predictions critically depend on the assumed density dependence of growth: when this is weak, life history evolution might be necessary to prevent stock collapse, whereas when a stronger density dependence estimated from recent data is used, the role of evolution in explaining fisheries-induced trait changes is diminished. Our integrative analysis of density-dependent growth, multitrait evolution, and stock-specific time series data underscores the importance of jointly considering evolutionary and ecological processes, enabling a more comprehensive perspective on empirically observed stock dynamics than previous studies could provide.

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On distributional responses of North Atlantic fish to climate change
  • Jan 1, 2005
  • ICES Journal of Marine Science
  • G.A Rose

Abstract Changes in fish distribution and climate in the North Atlantic have been observed for millennia by seafaring peoples, chronicled in many historical anecdotes, and recently studied systematically. For temperate to Arctic North Atlantic fish, a literature compendium of limits of temperature, salinity, and depth during feeding and spawning was used to investigate factors that influence distribution. Latitude and depth were negatively correlated with species number and density. Peak numbers of species feed at 0–4°C, but spawn at 2–7°C and salinities of 32.5–33.5. Principal components of feeding depths and temperatures suggested four groups of species: (i) small pelagics characterized by shallow habitat and cooler temperatures; (ii) most groundfish in deeper and warmer waters; (iii) warm-water large pelagics; and (iv) deepwater species. Spawning temperatures, salinities, depths, and timing produced groupings consistent with feeding components for pelagics, but differing for distant migrants such as tunas. Principal components (PCA) of spawning characteristics explained 56% of the variance in species resilience (doubling time), while PCA of feeding characteristics explained only 23%. We infer that the small pelagics capelin (Mallotus villosus) and herring (Clupea harengus) react strongly and quickly to climate change because of their physiological limits and potential for fast population growth. Verification comes from Icelandic and Greenland waters, which warmed considerably during 1920–1940, and where capelin, herring, cod (Gadus morhua), and other species shifted north very quickly.

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The influence of long tides on ecosystem dynamics in the Barents Sea
  • Dec 6, 2008
  • Deep Sea Research Part II: Topical Studies in Oceanography
  • Harald Yndestad

The influence of long tides on ecosystem dynamics in the Barents Sea

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Towards the Optimal Management of the Northeast Arctic Cod Fishery
  • Jun 17, 2011
  • SSRN Electronic Journal
  • Andries Richter + 3 more

Towards the Optimal Management of the Northeast Arctic Cod Fishery

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Long-term change in a behavioural trait: truncated spawning distribution and demography in Northeast Arctic cod
  • Dec 5, 2014
  • Global Change Biology
  • Anders Frugård Opdal + 1 more

Harvesting may be a potent driver of demographic change and contemporary evolution, which both may have great impacts on animal populations. Research has focused on changes in phenotypic traits that are easily quantifiable and for which time series exist, such as size, age, sex, or gonad size, whereas potential changes in behavioural traits have been under-studied. Here, we analyse potential drivers of long-term changes in a behavioural trait for the Northeast Arctic stock of Atlantic cod Gadus morhua, namely choice of spawning location. For 104 years (1866–1969), commercial catches were recorded annually and reported by county along the Norwegian coast. During this time period, spawning ground distribution has fluctuated with a trend towards more northerly spawning. Spawning location is analysed against a suite of explanatory factors including climate, fishing pressure, density dependence, and demography. We find that demography (age or age at maturation) had the highest explanatory power for variation in spawning location, while climate had a limited effect below statistical significance. As to potential mechanisms, some effects of climate may act through demography, and explanatory variables for demography may also have absorbed direct evolutionary change in migration distance for which proxies were unavailable. Despite these caveats, we argue that fishing mortality, either through demographic or evolutionary change, has served as an effective driver for changing spawning locations in cod, and that additional explanatory factors related to climate add no significant information.

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We evaluate a 54-y survey time series for the Delaware Bay oyster beds in New Jersey waters to identify the characteristics of regime shifts in oyster populations and the influence of MSX and Dermo diseases on population stability. Oyster abundance was high during the 1970s through 1985. Oyster abundance was low at the inception of the time series in 1953, remained low through 1969, and has been low since 1985 and very low since 2000. Natural mortality was low in most years prior to the appearance of MSX in 1957. From 1957 through 1966, natural mortality generally remained above 10% annually and twice exceeded 20%. Natural mortality remained well below 15% during the 1970s and into the early 1980s when oyster abundance was continuously high. The largest mortality event in the time series, an MSX epizootic that resulted in the death of 47% of the stock, occurred in 1985. Mortality rose again with the incursion of Dermo in 1990 and has remained above 15% for most years since that time and frequently has exceeded 20%. The primary impact of MSX and Dermo diseases has been to raise natural mortality and ultimately to cause a dispersed stock to retreat into its habitat of refuge in the moderately low salinity reach of the bay. The time series of oyster abundance on the New Jersey oyster beds of Delaware Bay is dominated by two regime shifts, the 1970 abundance increase that was maintained for about 15 y thereafter, and the 1985 abundance decrease that continues through today. These two regime shifts ushered in long-term periods of apparent constancy in population dynamics. The 1985 regime shift was induced by the largest MSX epizootic on record that produced high mortalities throughout a population distributed broadly throughout its habitat range after 15 y of high abundance. A putative new regime commenced circa 2000 as a consequence of a series of Dermo epizootics. Mortalities routinely exceeded 20% of the population annually during this period, with the consequence of a greater degree of stock consolidation than any previous time in the 54-y record. Extreme consolidation of the stock would appear to be a characteristic of the population's response to Dermo disease. The 1970-1984 and post-1985 regimes each were ushered in by a confluence of events unique in the 54-y time series. Each was characterized by a period of relative stability in population abundance. However, the stability in total population abundance belies a more dynamic process of stock redistribution during both time intervals, demonstrating that the appearance of constancy in stock abundance is not necessarily a result of invariant stock dynamics. Rather, the Delaware Bay oyster time series suggests that regime shifts delimit periods during which differential, often offsetting, local trends impart similar abundance levels, and thus constancy at the level of the stock masks substantive changes in local population dynamics potentially fostering future catastrophic changes in population-level attributes. Understanding such regime shifts will likely determine the success of decadal management goals more so than measures designed to influence population abundance.

  • Research Article
  • Cite Count Icon 43
  • 10.1111/cobi.12078
Effects of an Invasive Plant on Population Dynamics in Toads
  • May 21, 2013
  • Conservation Biology
  • Daniel A Greenberg + 1 more

When populations decline in response to unfavorable environmental change, the dynamics of their population growth shift. In populations that normally exhibit high levels of variation in recruitment and abundance, as do many amphibians, declines may be difficult to identify from natural fluctuations in abundance. However, the onset of declines may be evident from changes in population growth rate in sufficiently long time series of population data. With data from 23 years of study of a population of Fowler's toad (Anaxyrus [ = Bufo] fowleri) at Long Point, Ontario (1989-2011), we sought to identify such a shift in dynamics. We tested for trends in abundance to detect a change point in population dynamics and then tested among competing population models to identify associated intrinsic and extrinsic factors. The most informative models of population growth included terms for toad abundance and the extent of an invasive marsh plant, the common reed (Phragmites australis), throughout the toads' marshland breeding areas. Our results showed density-dependent growth in the toad population from 1989 through 2002. After 2002, however, we found progressive population decline in the toads associated with the spread of common reeds and consequent loss of toad breeding habitat. This resulted in reduced recruitment and population growth despite the lack of significant loss of adult habitat. Our results underscore the value of using long-term time series to identify shifts in population dynamics coincident with the advent of population decline.

  • Research Article
  • Cite Count Icon 4
  • 10.1071/mf18286
Predicting catch per unit effort from a multispecies commercial fishery in Port Phillip Bay, Australia
  • Sep 10, 2019
  • Marine and Freshwater Research
  • Karina L Ryan + 1 more

Quantitative models that predict stock abundance can inform stock assessments and adaptive management that allows for less stringent controls when abundance is high and environmental conditions are suitable, or tightening controls when abundance is low and environmental conditions are least suitable. Absolute estimates of stock abundance are difficult and expensive to obtain, but data from routine reporting in commercial fisheries logbooks can provide an indicator of stock status. Autoregressive integrated moving average (ARIMA) models were constructed using catch per unit effort (CPUE) from commercial fishing in Port Phillip Bay from 1978–79 to 2009–10. Univariate and multivariate models were compared for short-lived species (Sepioteuthis australis), and species represented by 1–2 year-classes (Sillaginodes punctatus) and 5–6 year-classes (Chrysophrys auratus). Simple transfer models incorporating environmental variables produced the best predictive models for all species. Multivariate ARIMA models are dependent on the availability of an appropriate time series of explanatory variables. This study demonstrates an application of time series methods to predict monthly CPUE that is relevant to fisheries for species that are short lived or vulnerable to fishing during short phases in their life history or where high intra-annual variation in stock abundance occurs through environmental variability.

  • Research Article
  • Cite Count Icon 9
  • 10.1023/a:1011087604758
The Influence of Hydroregime on Catch, Abundance And Recruitment of the Catfish, Chrysichthys Nigrodigitatus (Bagridae) and the Bonga, Ethmalosa Fimbriata (Clupeidae) of Southeastern Nigeria's Inshore Waters
  • May 1, 2001
  • Environmental Biology of Fishes
  • Boniface S Moses

This paper investigates the relationship between the hydroclimatic parameters (rainfall and flood index) and the catch, stock abundance and recruitment of the catfish, Chrysichthys nigrodigitatus (Bagridae) and the bonga, Ethmalosa fimbriata (Clupeidae) of southeastern Nigeria's inshore waters. For C. nigrodigitatus, most peaks in the mean biomass \((\Delta \% {\bar B})\) and recruitment \((\Delta \% {\bar R(n)})\) curves occurred during the 'wet' years, i.e., years for which the percentage deviations of rainfall \((\Delta \% \overline {{RF}} )\) and flood index \((\Delta \% \overline {{FI}} - 1)\) from their means remained above their averages. Catch and abundance respectively showed good positive linear correlation with the flood index. E. fimbriata behaved differently; some peaks in the \(\Delta \% {\bar B}\) and \(\Delta \% {\bar R(n)}\) curves occurred in the 'wet' and some in the 'dry' years; and there was no correlation between the annual catch of bonga and either the rainfall or the flood index. The hypothesis, that linear relationships exist between the interannual variations in the hydroregime and the yearly fluctuations in the catch and population structure of some coastal and estuarine fishes, holds true for the catfish, C. nigrodigitatus, but not for bonga, E. fimbriata.

  • Research Article
  • Cite Count Icon 2
  • 10.1086/mre.5.4.42628938
Fixed or Variable Catch Quotas? The Importance of Population Dynamics and Stock Dependent Costs
  • Jan 1, 1988
  • Marine Resource Economics
  • Rögnvaldur Hannesson

This article considers two questions concerning fluctuations in fish stocks and catches: Are they possibly beneficial and would fishery managers opt for a strategy that enchances stock or catch quota variability, or both? For this analysis, three models are followed: in the first, variations in stock abundance are stochastic with no population dynamics; in the second, the stock in a certain time period depends on a stochastic recruitment to the stock and survivors from the previous time period; and the third is a year class model that has more complicated population dynamics but is more realistic.

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