Changes in spawning stock structure strengthen the link between climate and recruitment in a heavily fished cod (Gadus morhua) stock

Atlantic cod (Gadus morhua) is one of the commercially most important fish species in the North Atlantic and plays a central role in several ecosystems. Fishing pressure has been heavy over a prolonged period and the recent decades have shown dramatic decline in abundance of many stocks. The Arcto‐Norwegian (or North‐east Arctic) cod stock in the Barents Sea is now the largest stock of Atlantic cod. Recruitment to this stock has varied extensively during the last 60 yr. There is evidence for fluctuations in climate, particularly sea temperature, being a main cause for this variability, higher temperatures being favourable for survival throughout the critical early life stages. Our studies of time series present compelling evidence for a strengthening of the climate–cod recruitment link during the last decades. We suggest this is an effect of the age and length composition of the spawning stock having changed distinctly. The age of the average spawner has decreased by more than 3 yr from between 10 and 11 in the late 1940s to 7–8 in the 1990s, average length from just above 90 cm to around 80 cm. The number of age classes contributing to the spawning stock has also decreased, while the number of length groups present increased slightly. Significant decrease in age of spawners has frequently been described for other heavily fished stocks worldwide. We therefore find it likely that the proposed mechanism of increased influence of climate on recruitment through changes in the spawning stock age and size composition is of a general nature and might be found in other systems.     

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

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.     

Greenland cod (Gadus morhua): modeling recruitment variation during the second half of the 20th century

Two approaches were used to qualify observed variability in Greenland cod (Gadus morhua) recruitment. In the first analysis, we used the linear trend of the Greenland cod recruitment time series and climatic variables, such as air temperatures from the Denmark Strait and wind conditions off East Greenland and Southwest Greenland, to explain the interannual variation in cod recruitment off Greenland. The model resulting from this ‘trend/environmental approach’, explained 79% of the interannual variation in cod recruitment off Greenland. In the second, analytical approach, the ‘regime approach’, multiple linear regression models were used, with the input data being the time series of cod recruitment and spawning stock biomass (SSB) from Iceland and Greenland, sea surface and air temperatures around Greenland, and zonal wind components between Iceland and Greenland. Model results indicated that, during the decades between 1950 and 1990, there were three different cause–effect regimes which significantly influenced the variability of cod recruitment. The three regimes included: (a) the 1950s and 1960s, a regime with favorable sea surface temperatures and a self‐sustaining cod stock off Greenland with high SSB that produced a series of above‐average, strong year classes; (b) the 1970s and 1980s, a regime of declining SSB and recruitment, with recruitment dependent on advection from Iceland; and (c) the 1990s, when the advective potential for recruitment from the Icelandic cod stock was the only available source for replenishment of the Greenland cod stocks, because cod recruitment in Greenland waters was negligible. The three models explained 76–77% of the observed interannual variation in cod recruitment off Greenland. Both approaches suggested that advective factors were the dominant influences for cod recruitment in the ‘Iceland–Greenland System’.     

Match/mismatch predictions of spawning duration versus recruitment variability

According to the match/mismatch hypothesis, larval fish survival and eventual recruitment is dependent on the offset time between the peaks of abundance of larvae and their planktonic prey. A rudimentary larval food supply model is developed to determine the dependence of food availability on the mismatch between peaks. The model predicts that recruitment variability should increase as spawning duration decreases, a result which is moderately supported by an analysis of Atlantic cod (Gadus morhua) data.     

Stability of water masses-impact on cod recruitment off West Greenland

Based upon the hypothesis of Meyer (1968: Hansa, 16:2–4) that periods of low stability of the water column in autumn should parallel good cod year-classes in the following year, the paper analyzes available time series of cod recruitment, subsurface oceanographic stability data and wind stress data off West Greenland. No significant correlations were found between stability, wind stress and recruitment when considering the entire. length of available time series. However, splitting the autumn-based oceanographic data set into a pre- 1970s part (when warm climatic conditions and high recruitment prevailed), and a recent part (for the past 20 years of cooling climate) yields significant coherence of recruitment and environmental conditions for the cold climate period. For the warm climate period, only occasional consistency is shown. The possible influence of the Great Salinity Anomaly on recruitment of cod is discussed. It is suggested that during the past 20 years a new correlation mechanism, an advective coupling instead of low stability coupling, may explain the variability of recruitment.     

Influence of larval transport and temperature on recruitment dynamics of North Sea cod (Gadus morhua) across spatial scales of observation

The survival of fish eggs and larvae, and therefore recruitment success, can be critically affected by transport in ocean currents. Combining a model of early‐life stage dispersal with statistical stock–recruitment models, we investigated the role of larval transport for recruitment variability across spatial scales for the population complex of North Sea cod (Gadus morhua). By using a coupled physical–biological model, we estimated the egg and larval transport over a 44‐year period. The oceanographic component of the model, capable of capturing the interannual variability of temperature and ocean current patterns, was coupled to the biological component, an individual‐based model (IBM) that simulated the cod eggs and larvae development and mortality. This study proposes a novel method to account for larval transport and success in stock–recruitment models: weighting the spawning stock biomass by retention rate and, in the case of multiple populations, their connectivity. Our method provides an estimate of the stock biomass contributing to recruitment and the effect of larval transport on recruitment variability. Our results indicate an effect, albeit small, in some populations at the local level. Including transport anomaly as an environmental covariate in traditional stock–recruitment models in turn captures recruitment variability at larger scales. Our study aims to quantify the role of larval transport for recruitment across spatial scales, and disentangle the roles of temperature and larval transport on effective connectivity between populations, thus informing about the potential impacts of climate change on the cod population structure in the North Sea.     

Linking cod (Gadus morhua) and climate: investigating variability in Irish Sea cod recruitment

How climatic variability and anthropogenic pressures interact to influence recruitment is a key factor in achieving sustainable resource management. However, the combined effects of these pressures can make it difficult to detect non‐stationary interactions or shifts in the relationships with recruitment. Here we examine the links between climate and Irish Sea cod recruitment during a period of declining spawning stock biomass (SSB). Specifically, we test for a shift in the relationship between recruitment, SSB and climate by comparing an additive (generalized additive model, GAM) and non‐additive threshold model (TGAM). The relationship between recruitment success, SSB and the climatic driver, sea surface temperature, was best described by the TGAM, with a threshold identified between recruitment and SSB at approximately 7900 t. The analysis suggests a threshold shift in the relationship between recruitment and SSB in Irish Sea cod, with cod recruitment being more sensitive to climatic variability during the recent low SSB regime.     

Effects of dietary protein and lipid level, and water temperature, on the post-feeding oxygen consumption of Atlantic cod and haddock

Tank respirometry was used to study the interactive effects of protein:lipid level (55%:11% vs. 42%:16%; both diets isoenergetic) and temperature (11, 6 and 2 °C) on the magnitude and duration of specific dynamic action (SDA) in juvenile Atlantic cod and haddock. The protein:lipid level did not affect any measured variable. However, numerous temperature and species effects were observed. For example, although the maximum post-feeding oxygen consumption (30–50% above routine metabolic rate; RMR) and SDA duration (∼55–85 h; SDA_DUR) were not affected by temperature, SDA_DUR g⁻¹ of food increased from 11 to 2 °C (from ∼3 to 12 h g food⁻¹). While absolute SDA (mg O₂) decreased by ∼60–65% in cod and ∼75% in haddock from 11 to 2 °C, due to a concomitant decrease in food consumption from ∼2.0% to 0.6% body mass, SDA comprised between 3.3% and 5.2% of the dietary energy content at all temperatures. Finally, RMR at 11 and 2 °C and SDA_DUR at 2 °C were 25–35% and 25% greater in cod, respectively, as compared with haddock. These results suggest that feeding reduced protein diets at low water temperatures is unlikely to improve the growth of these species.     

Juvenile cod (Gadus morhua) mortality and the importance of bottom sediment type to recruitment on Georges Bank

The demersal settlement of pelagic juvenile fish has been considered a critical period in which the final adjustment is made to the size of a year class. Distribution patterns of pelagic and recently settled juvenile cod (Gadus morhua) were examined from nine surveys on Georges Bank during the summer over 5 years, 1984–1989, to relate juvenile survival to the sedimentary environment. Pelagic juveniles were widespread across Georges Bank in June, and by mid‐July they occurred on all bottom types from sand to gravel on eastern Georges Bank. However, by late July‐early August they were mostly abundant on the northeastern edge gravel deposit, which with its complex relief, provides abundant prey and refuge from predators. A bank‐wide estimate of pelagic juvenile abundance in 1986 and 1987 was used to assess mortality of the recently settled juveniles and to evaluate the relative importance of survivors from the northeastern edge gravel area to recruitment of the Georges Bank population. Settlement mortality rates over 1–2 months on the northeastern gravel area ranged from 3 to 8% day^?1, which compared reasonably with other studies. The seasonal abundance of the pelagic juveniles was almost an order of magnitude higher in 1987 than 1986; however, recruitment at age 1 was similar, indicating that a high mortality of the demersal juveniles occurred in 1987. The limited northeastern gravel area on Georges Bank may represent a survival bottleneck depending on the variability in the distribution and abundance of juvenile cod settlement in relation to that of their predators.     

Weight-based yield per recruitment and spawning-biomass per recruitment analyses of pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> off the Pacific coast of southern Hokkaido,Japan

ABSTRACT:   The present study assessed the stock state of Pacific cod Gadus macrocephalus caught off the coast of southern Hokkaido, Japan. Weight-based yield per recruitment (YPR) and spawning-biomass per recruitment (SPR) analyses were used for this assessment. The current fishing mortality (average from 1998 to 2000) was 0.65 and weight at first capture was 0.5 kg bodyweight. Under these fishing pressures, the YPR of Pacific cod in southern Hokkaido was 1.06 kg/recruitment and percentage of SPR (%SPR) was 6.9%. The %SPR was lower than the critical limit at 20%SPR. The main reason that values of both YPR and %SPR were not optimum, would be that the weight at first capture was too small. Raising the weight at first capture was thought to be a better strategy from the biological viewpoint, and reducing fishing mortality to 0.3 would be the next alternative strategy from the fisheries management viewpoint.     

Temperature influences on growth of unfished juvenile Northern cod (Gadus morhua) during stock collapse

This study examined growth of unfished juvenile Northern cod (Gadus morhua) off Newfoundland concomitant with stock collapse in the cold early 1990s. Two unpublished data sets were examined, one from collapse‐period trapping sites along the northeast coast of Newfoundland and one from a post‐collapse inshore trawl survey. Cumulative surface and bottom temperatures were significant predictors of growth rates of the young fish with year‐classes born during collapse experiencing slower growth than those born during subsequent warming. Relationships between accrued temperature and growth were consistent across periods, with slow growth of collapse‐period fish reflecting slower accumulation of temperature‐at‐age. Temperature influences were spatially broad‐based with no significant differences in growth rates for fish captured along the entire northeast coast of Newfoundland. Predicted differences in growth rates for collapse versus recovery year‐classes were proportional to cumulative surface temperatures but not cumulative bottom temperatures. Although significant, temperature effects on growth were relatively unimportant at youngest ages. Overall, growth differences between periods were small but large differences occurred between slowest and fastest growing year‐classes. The results suggest initial responses to increasing temperatures were delayed following collapse. We conclude that although temperature was an important determinant of dampened productivity that it alone cannot account for the collapse and slow recovery of the stock. This is the first known study to directly quantitatively link temperature impacts to an unfished component of the Northern cod stock complex during collapse, removing need for implicit assumptions about whether or not cold conditions contributed to the collapse of this iconic fish stock.     

Incorporating an environmental stock–recruitment relationship in the assessment of Pacific cod (Gadus macrocephalus)

The robustness of a previously described environmentally mediated stock–recruitment relationship for Pacific cod in Hecate Strait, BC, Canada was tested with 10 yr of additional data. The original analysis tested several alternative hypotheses and concluded that water transport through Hecate Strait, as indicated by sea level height, coupled with cod spawning biomass formed the best model. The present analysis indicates the relationship held through the 1990s. The implications of variation in sea level on stock production were investigated with a delay‐difference stock production model that included an environmentally mediated stock–recruitment relationship. The model predicted that the maximum fishery yield would vary between 1750 and 3670 t yr^?1 over the observed range of sea level height, and the estimated unsustainable fishing mortality during periods of low productivity would be only 0.5 times that in periods of high productivity.     

东平湖刀鲚生物学参数、单位补充量渔获量及产卵亲体量评估

为完善东平湖刀鲚(Coilia nasus)的基础生物学数据, 并为实现其资源的可持续利用提供技术指导和理论依据, 本研究根据 2021—2023 年东平湖定置三层多目刺网调查获得的刀鲚体长、体重数据, 利用 ELEFAN_GA 技术对刀鲚的生长和死亡参数进行估算, 拟合 Von Bertalanffy (VBGF)生长方程, 构建了基于体长结构的单位补充量渔获量(YPR)和单位补充量产卵亲体量(SSBPR)模型。结果显示, 东平湖刀鲚体长范围为 4.9~26.2 cm, 平均体长为 (12.2±2.4) cm, 体重范围 1.1~51.8 g, 平均体重为(6.6±4.5) g; 使用基于 bootstrap 的 ELEFAN_GA 方法估算刀鲚渐进体长 L_∞=31.65 cm, 生长速率 K=0.66, 理论生产起点年龄 t₀= ‒0.23, 季节性生产振幅 C=0.65, 1 年中生产最快的时间段 t_s=0.40; 刀鲚的总死亡系数 Z=2.71, 自然死亡系数 M=1.074, 捕捞死亡系数 F=1.641、开发率 E=0.604, 平均选择体长 L₅₀=9.15 cm; YPR 模型结果显示, 随着 F 增大, YPR 值呈现先上升后下降的趋势, 生物学参考点 F_0.1 和 F_max 的值分别为 0.978 和 2.204; SSBPR 模型结果显示随着捕捞强度的增大, SSBPR 呈下降趋势, 现阶段捕捞强度介于 F_20%(2.848)与 F_40%(1.065)之间, 略大于 F_35%(1.286)。综上所述, 东平湖刀鲚生长表现出明显的季节性变化规律, 在与其他地理群体对比中, 东平湖刀鲚在生物学参数上的差异体现了其生长过程的空间异质性, K 值最大表明其可以用最短的时间生长到接近极限体长, 说明东平湖饵料丰富, 而同时较高的开发率和较低的开捕体长则表明目前东平湖刀鲚已处于过度开发状态, SSBPR 曲线得出现阶段捕捞强度小于极限值 F_20%, 略大于生物学参考点 F_35%。研究结果表明, 为使东平湖刀鲚资源达到生态、经济效益平衡, 可在适当减少渔船数量以降低捕捞强度的情况下, 通过增大网目尺寸以增加开捕体长来保持其可持续发展。     

Environmental influences on the recruitment of Newfoundland/Labrador Cod*

This paper investigates possible linkages between the environment and year-class success of Newfoundland/ Labrador cod. Aspects of the physical environment relevant to recruitment are reviewed and the current knowledge of the patterns of spawning and larval occurrence is summarized. A critical review of investigations aimed at correlating recruitment with climactic fluctuations is made and concludes that these studies have been largely fruitless. A series of numerical simulations of particle drift are presented to investigate the potential importance of storms and to try to identify nursery grounds. These simulations suggest that Labrador Shelf eggs and larvae are segregated from Grand Banks eggs and larvae. Favorable storm tracks appear to be required for significant numbers of eggs and larvae to reach the bays of Northeast Newfoundland.     

Oceanographic influences on the distribution of Baltic cod, Gadus morhua, during spawning in the Bornholm Basin of the Baltic Sea

The Baltic Sea is a stratified, semienclosed sea typified by a low-salinity surface layer and a deep saline layer of varying volume, salinity, temperature and oxygen concentration. The relationships between these oceanographic factors and the distribution of Baltic cod are presented, utilizing results from a survey carried out during the 1995 spawning period in the Bornholm Basin, at present the main spawning area of this stock. Cod distribution, abundance and population structure were estimated from hydroacoustic and trawl data and related to hydrographic parameters as well as to bottom depth. In the central basin, cod were aggregated in an intermediate layer about 15 m thick. This area of peak abundance was defined at its upper limit by the halocline and at the lower limit by oxygen content. The majority of individuals caught in the basin centre were in spawning or pre-spawning condition with a high proportion of males to females. On the basin slopes, aggregations of cod were found near the bottom. These individuals were mainly immature and maturing stages with an increasing proportion of females to males with size. Salinity and oxygen conditions were found to be the major factors influencing the vertical and horizontal distribution of adult cod. Abundance of immature cod was also positively related to decreasing bottom depths. The effect of temperature was minor. The observed size- and sex-dependent spawning aggregation patterns, in association with habitat volume and stock size, may influence cod catchability and thereby the assessment and exploitation patterns of this stock.     

Influence of incubation temperature on body movements of Atlantic cod (Gadus morhua L.) embryos and on size at hatch

Body movements of cod (Gadus morhua L.) embryos reared from fertilization to hatch at 5.4°C were observed at various stages of development and at six experimental temperatures ranging from 0–10°C. Frequency of cod embryo body movements increased from zero at 42 degree‐days post fertilization to maximal at 73–82 degree‐days (1 or 2 days prior to hatch). Embryos were most active at 2°C (mean of 5.5 movements per 10 min), with activity declining to less than 1/10 min at 8–10°C. Lengths of hatched cod larvae reared at a series of constant temperatures (from 4–10°C) from fertilization to hatch were greater at lower incubation temperatures. Incubation temperatures of 2–4°C were found to be optimal for incubation of cod eggs.     

Relationships between temperature and landing patterns in the southern Labrador and eastern Newfoundland inshore cod fishery

The temporal pattern of cod landings in the inshore trap fishery off the east coast of Newfoundland and Labrador is examined in this study. Unlike previous studies which have been broad scale or of a localized nature, this analysis looks at variations within fishing seasons and among fishing areas in trap landings. A significant relationship between water temperature and the weekly pattern is demonstrated. Significant spatial patterns in weekly landings are described and related to longshore differences in the seasonal variation in temperature.     

Effects of different step‐wise temperature increment regimes during egg incubation of Atlantic cod (Gadus morhua L.) on egg viability and newly hatched larval quality

We carried out an experiment to determine how rapidly the early incubation temperature of Atlantic cod eggs can be increased without affecting normal embryonic development and hatching. Atlantic cod eggs were incubated at a constant low temperature (4.5 ± 0.5°C; T5 – control) and four temperature increment treatments where the temperatures were increased stepwise from 4.5°C at zygote stage to 9.5 ± 05°C (T1‐8 h, T2‐32 h, T3‐64 h and T4‐96 h). Embryonic cell symmetry, embryonic mortality, hatching success and larval skeletal abnormalities, length and yolk sac volume were recorded. Larval samples were also taken at hatch for histological analysis. Except for higher egg mortality and lower hatching success in the T1, the differences among experimental groups were minor. Cell asymmetries and embryo mortalities were not significantly different between the control and T2–T4 treatment groups. Control larvae were significantly longer and had smaller yolk reserves at hatch than T1–T4 larvae and larvae from T2 had the largest yolk reserves. Tissue and organ histology of hatched larvae were similar. Considering embryonic cleavage pattern, hatching success and larval morphology and histology, a gradual increment of temperature in 32 h seems to be the better choice for future developmental programming studies in Atlantic cod.     

Re‐evaluating the effect of wind on recruitment in Gulf of Maine Atlantic Cod (Gadus morhua) using an environmentally‐explicit stock recruitment model

A previous study documented a correlation between Atlantic Cod (Gadus morhua) recruitment in the Gulf of Maine and an annual index of the north component of May winds. This correlation was supported by modeling studies that indicated strong recruitment of Gulf of Maine Atlantic Cod results from high retention of spring‐spawned larvae in years when winds were predominately out of the north, which favor downwelling. We re‐evaluated this relationship using updated recruitment estimates and found that the correlation decreased between recruitment and wind. The original relationship was largely driven by two recruitment estimates, one of which (2005 year‐class) was highly uncertain because it was near the terminal year of the assessment. With additional data, the updated assessment estimated lower recruitment for the 2005 year‐class, which consequently lowered the correlation between recruitment and wind. We then investigated whether an environmentally‐explicit stock recruit function that incorporated an annual wind index was supported by either the original or updated assessment output. Although incorporation of the annual wind index produced a better fitting model, the uncertainty in the estimated parameters and the implied unexploited conditions were not appropriate for providing management advice. These results suggest the need for caution in the development of environmentally‐explicit stock recruitment relationships, in particular when basing relationships and hypotheses on recruitment estimates from the terminal years of stock assessment models. More broadly, this study highlights a number of sources of uncertainty that should be considered when analyzes are performed on the output of stock assessment models.