Reproductive potential of the eastern baltic cod <Emphasis Type="Italic">Gadus morhua callarias</Emphasis> L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997–2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

Evolutionary response to size-selective mortality in an exploited fish population

Many collapsed fish populations have failed to recover after a decade or more with little fishing. This may reflect evolutionary change in response to the highly selective mortality imposed by fisheries. Recent experimental work has demonstrated a rapid genetic change in growth rate in response to size-selective harvesting of laboratory fish populations. Here, we use a 30-year time-series of back-calculated lengths-at-age to test for a genetic response to size-selective mortality in the wild in a heavily exploited population of Atlantic cod (Gadus morhua). Controlling for the effects of density- and temperature-dependent growth, the change in mean length of 4-year-old cod between offspring and their parental cohorts was positively correlated with the estimated selection differential experienced by the parental cohorts between this age and spawning. This result supports the hypothesis that there have been genetic changes in growth in this population in response to size-selective fishing. Such changes may account for the continued small size-at-age in this population despite good conditions for growth and little fishing for over a decade. This study highlights the need for management regimes that take into account the evolutionary consequences of fishing.     

Allele frequency covariance among cohorts and its use in estimating effective size of age-structured populations

A general expression for the covariance of allele frequencies among cohorts in age-structured populations is derived. The expression is used to extend the so-called temporal method for estimating effective population size from allele frequency shifts among samples from cohorts born any number of years apart. Computer simulations are used to check on the accuracy and precision of the method, and an application to coastal Atlantic cod is presented.     

Effect of size-selective mortality on growth of coastal cod illustrated by tagging data and an individual-based growth and mortality model

In a release experiment with cod in Norway, the apparent mean growth rates of 3+ cod, calculated by sampling the released cohorts at different ages, were very slow (<0·08 mm day⁻¹). However, when individual growth rates of individual tagged cod of the same size range were measured, the mean growth rates were much faster (0·24 mm day⁻¹). These observations were attributed to size-selective fishing mortality and were illustrated by an individual based simulation model of a cohort of cod with variable individual growth rates. The effects on mean length at age of the surviving cohort of increasing fishing intensity were demonstrated. The model showed that size-selective fishing with the observed individual growth variation, removed the fastest-growing individuals at proportionally higher rates than the slower-growing ones, leading to decreased apparent mean growth rate. The fishing pattern which gave the optimum yield, changed when individual variation was included, and when the apparent growth rate was used in the model the yield per recruit reduced dramatically. This study has shown that individual growth heterogeneity and size-selective mortality are factors which should be considered in future fisheries management models.     

Comparison of lethal and non-lethal age-based growth estimation methodologies to assess an endemic bay population of Atlantic cod (Gadus morhua)

The Gilbert Bay, Labrador, MPA (Marine Protected Area) was created in 2005 to protect a genomically distinct and locally adapted bay resident population of Atlantic cod (Gadus morhua) from commercial overfishing. Since then research showed that significant numbers of individuals migrate beyond MPA boundaries creating a concern that fishing continues to impact this endemic population. As part of research and MPA monitoring to assess the health of the protected cod population, lethal sampling of Gilbert Bay cod was conducted from 1998 to 2009 to gather length-at-age data from otoliths. Since then only non-lethal sampling methods were used so as to help conserve the protected population. This study was conducted to compare non-lethally (mark-recapture and length-frequency) obtained age and growth data with lethally (otolith-length-at-age) obtained data. The three methods produced similar results for the 1998 to 2009 period, and both of the non-lethal methods considered were consistent over the 20 year study, confirming their preference over lethal methods for this small endemic fish population.     

Early Environmental Influences on Growth of Arcto-Norwegian Cod, Gadus Morhua, From The 0-group To Adults

A high growth rate for Arcto-Norwegian cod, Gadus morhua, in the Barents Sea and adjacent areas from the larva period to the 0-group enhances survival and ultimately recruitment to the fishery. However, it appeared that high growth rates for a cohort through the 0-group were not continued as the cohort ages. Based on survey data, there was a significant negative correlation between the average length at the 0-group and its average length at ages 2 through 8. We provided evidence suggesting that this phenomenon was caused by the inter-annual variability in inflow of warm, prey-rich Atlantic water into the Barents Sea from the Norwegian Sea. Enhanced inflow provided favorable conditions for cod growth during the larva and juvenile pelagic intervals. However, this same strong inflow carried a proportion of the cohort farther to the east in the Barents Sea, where the bottom water is colder than in the west. The colder conditions experienced by such cohorts, as compared to cohorts that have a more westerly settlement, led to slower growth prior to age 2. Slow growth during this interval appeared to be the reason for these cohorts' relatively smaller mean length at older ages.     

Bay-scale population structure in coastal Atlantic cod in Labrador and Newfoundland, Canada

Polymorphisms at five microsatellite DNA loci provide evidence that Atlantic cod Gadus morhua inhabiting Gilbert Bay, Labrador are genetically distinguishable from offshore cod on the north-east Newfoundland shelf and from inshore cod in Trinity Bay, Newfoundland. Antifreeze activity in the blood suggests that Gilbert Bay cod overwinter within the Bay. Gilbert Bay cod are also smaller (weight and length) for their age and consequently less fecund for their age, than cod elsewhere within the northern cod complex. The productivity and recruitment potential of coastal cod off Labrador may thus be much lower than that of offshore northern cod or of inshore cod farther south, implying that a more conservative management strategy may be required for cod from coastal Labrador than traditionally practised for northern cod inhabiting less harsh environments. Relatively high F _ST and R _ST measures of population structure suggest that important barriers to gene flow exist among five components that include two inshore (Gilbert and Trinity Bay) and three offshore cod aggregations on the north-east Newfoundland Shelf and the Grand Bank. D _A and D _SW estimates of genetic distance that involve Gilbert Bay cod are approximately three- and 10–fold larger, respectively, than estimates not involving Gilbert Bay cod. The differences between inshore cod from Gilbert Bay and Trinity Bay raise the possibility that other genetically distinguishable coastal populations may exist, or may have existed prior to the northern cod fishery collapse. Harvesting strategies for northern cod should recognize the existence of genetic diversity between inshore and offshore components as well as among coastal components.     

Oxygen and salinity characteristics of predator–prey distributional overlaps shown by predatory Baltic cod during spawning

In the distributional overlap volume of Baltic cod Gadus morhua and its prey, studied in the Bornholm Basin in the southern Baltic Sea, only a fraction of the sprat Sprattus sprattus population vertically overlapped with the Baltic cod population. Sprat occurred in the intermediate water, in the halocline and in the bottom water, while herring Clupea harengus and Baltic cod occurred exclusively in the halocline and in the bottom water. Only parts of the sprat population were hence accessible for Baltic cod, and only a fraction of the sprat had access to the Baltic cod eggs below the halocline. Baltic cod–clupeid overlap volumes appeared to be determined by salinity stratification and oxygenation of the bottom water. Hydrography time series were used to estimate average habitat volumes and overlap from July to September in 1958–1999. In the 1999 survey spawning Baltic cod had greater ratios of empty stomachs and lower average rations than non-spawning Baltic cod. The average ration for Baltic cod caught within 11· 4 m from the bottom (demersal) did not differ from the average ration of Baltic cod caught in shallower waters (pelagic), because spawning and non-spawning Baltic cod in both strata were caught at equal rates. The diet of the Baltic cod caught demersally contained more benthic invertebrates, especially Saduria entomon, but Baltic cod caught pelagically also had fresh benthic food in their stomachs, indicating vertical migration of individual fish.     

A snapshot of cannibalism in 0-group Atlantic cod (Gadus morhua) in a marine pond

Cannibalism in 0-group Atlantic cod ( Gadus morhua ) was investigated in a semi-natural marine pond using stomach content analysis. About 6000 individuals were eradicated by rotenone treatment of the pond in November 1991. Total lengths of cod ranged from 7.4–28.8 cm with a median of 10.4 cm. The proportion of cannibalistic cod in the population was estimated at 1.3%. Cannibalism accounted for a daily loss of 1.1% of the population. The frequency of cod with ≥ 2 cod prey per stomach was higher in the population compared with a Poisson model. Predator-prey length ratios ranged between 1.6 and 3.2 when estimated from otolith-length relationships. Chesson's 1983 α-index indicated that cannibalistic cod had a positive selectivity for conspecifics < 9.0 cm in total length.     

Historic changes in length distributions of three Baltic cod (Gadus morhua) stocks: Evidence of growth retardation

Understanding how combinations of fishing effort and selectivity affect productivity is central to fisheries research. We investigate the roles of fishing regulation in comparison with ecosystem status for Baltic Sea cod stock productivity, growth performance, and population stability. This case study is interesting because three cod populations with different exploitation patterns and stock status are located in three adjacent but partially, ecologically different areas. In assessing stock status, growth, and productivity, we use survey information and rather basic stock parameters without relying on age readings. Because there is an urgent interest of better understanding of the current development of the Eastern Baltic cod stock, we argue that our approach represents partly a novel way of interpreting monitoring information together with catch data in a simplified yet more informative way. Our study reports how the Eastern and Western Baltic cod have gone toward more truncated size structures between 1991 and 2016, in particular for the Eastern Baltic cod, whereas the Öresund cod show no trend. We suggest that selective fishing may disrupt fish population dynamic stability and that lower natural productivity might amplify the effects of selective fishing. In support of earlier findings on a density‐dependent growth of Eastern Baltic cod, management is advised to acknowledge that sustainable exploitation levels for Eastern Baltic cod are much more limited than perceived in regular assessments. Of more general importance, our results emphasize the need to embrace a more realistic view on what ecosystems can produce regarding tractable fish biomass to facilitate a more ecosystem‐based fisheries management.     

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.     

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

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

Faster or slower: has growth of eastern Baltic cod changed?

Recent environmental changes have influenced the ecology and biology of eastern Baltic cod. Declining somatic condition, maturation at smaller size and restricted size distribution of the population suggest that growth rates have decreased between the early 2000s and the 2010s. Extensive age estimation problems have until now precluded testing of this hypothesis. This study presents evidence for a decrease in somatic growth rate of Baltic cod. Temporal patterns of growth, condition and maturation were analysed based on two complementary analyses: length frequency mode progression derived from DATRAS bottom trawl survey data and known-age samples, where size at age was back-calculated from daily otolith growth patterns. In the known-age samples, growth was positively related to somatic condition at capture with maturity dependent differences. Immature individuals had experienced significantly lower growth and were in lower condition at capture than mature individuals. Growth rates in the known-age samples were estimated at 9.5, 7.8 and 5.7?cm per year for age classes 1, 2 and 3 respectively. Growth between age 2 and 3 decreased significantly from 8.8?cm in the 1997 year class to 7.6?cm in the 2010 year class. While the 2001 and 2004 known-age samples were representative for the population, the 2013 sample was biased towards individuals with a higher condition and growth. Complementary length frequency analysis following the length mode of fish from age 2 to age 3 confirmed growth estimates from the early 2000s, while suggesting a 37.5% lower growth in 2013 compared with 2005.     

Allozyme analyses of the genus Trisopterus: taxonomic status and population structure of the poor cod

Genetic analysis of the four Trisopterus (Gadidae) taxa suggests that the interrelationships of the two morphs of poor cod ( T. minutus minutus in the Atlantic and T. minutus capelanus in the Mediterranean) should be reconsidered. The Mediterranean poor cod T. m. capelanus is more closely related to bib T. luscus than to the Atlantic poor cod, so the population structure in the Atlantic and Mediterranean poor cod must be considered separately. Among 635 Atlantic individuals there was some evidence of poor cod population differentiation (allele frequency heterogeneity test P <0·0005; F _ST=0·0135, P ≤0·0005). Levels of genetic variation were similar to those reported for related gadoid species. Some differentiation was present on the Norwegian coast (samples from Trondheimsfjord) and between the Faeroe Islands (Faeroe Bank) and the adjacent European coastal location. In contrast no statistically significant population differentiation was evident in Mediterranean poor cod, but fewer samples and individuals were screened.     

Historical DNA reveals the demographic history of Atlantic cod (Gadus morhua) in medieval and early modern Iceland

Atlantic cod (Gadus morhua) vertebrae from archaeological sites were used to study the history of the Icelandic Atlantic cod population in the time period of 1500–1990. Specifically, we used coalescence modelling to estimate population size and fluctuations from the sequence diversity at the cytochrome b (cytb) and Pantophysin I (PanI) loci. The models are consistent with an expanding population during the warm medieval period, large historical effective population size (N_E), a marked bottleneck event at 1400–1500 and a decrease in N_E in early modern times. The model results are corroborated by the reduction of haplotype and nucleotide variation over time and pairwise population distance as a significant portion of nucleotide variation partitioned across the 1550 time mark. The mean age of the historical fished stock is high in medieval times with a truncation in age in early modern times. The population size crash coincides with a period of known cooling in the North Atlantic, and we conclude that the collapse may be related to climate or climate-induced ecosystem change.     

Climate effects on size‐at‐age: growth in warming waters compensates for earlier maturity in an exploited marine fish

Over the past 3 decades, North Sea Atlantic cod (Gadus morhua) have exhibited variable length‐at‐age along with declines in spawning stock biomass and timing of maturity. Multiple factors affecting growth and development in fish acted on this economically important stock over the same period including warming waters and an intensive fishery. Here, we employ North Sea cod as a model population, exploring how a physiologically relevant temperature metric (the growing degree‐day, GDD; °C day) can be used to compare year‐classes on a physiologically relevant time‐scale, disentangling influences of climate (thermal history) on observed length‐at‐age trends. We conclude that the trends in North Sea cod length‐at‐age observed during the last three decades can be explained by a combination of temperature‐dependent growth increases and a trend toward earlier maturation, the latter likely induced by the intensive fishing pressure, and possibly evidence of fisheries‐induced evolution.     

Dynamics of coastal cod populations: intra- and intercohort density dependence and stochastic processes

Skagerrak populations of Atlantic cod (Gadus morhua L.) have been surveyed at several fixed stations since 1919. These coastal populations consist of local stocks with a low age of maturity and a short life span. We investigated 60 time-series of 0-group juveniles (i.e. young of the year) sampled annually from 1945 to 1994. An age-structured model was developed which incorporates asymmetrical interactions between the juvenile cohorts (0-group and 1-group; i.e. one-year-old juveniles) and stochastic reproduction. The model was expressed in delay coordinates in order to estimate model parameters directly from the time-series and thereby test the model predictions. The autocovariance structure of the time-series was consistent with the delay coordinates model superimposed upon a long-term trend. The model illustrates how both regulatory (density-dependent) and disruptive (stochastic) forces are crucial in shaping the dynamics of the coastal cod populations. The age-structured life cycle acts to resonance the stochasticity inherent in the recruitment process.     

Iteroparous reproduction strategies and population dynamics

Asymptotic relationships between a class of continuous partial differential equation population models and a class of discrete matrix equations are derived for iteroparous populations. First, the governing equations are presented for the dynamics of an individual with juvenile and adult life stages. The organisms reproduce after maturation, as determined by the juvenile period, and at specific equidistant ages, which are determined by the iteroparous reproductive period. A discrete population matrix model is constructed that utilizes the reproductive information and a density-dependent mortality function. Mortality in the period between two reproductive events is assumed to be a continuous process where the death rate for the adults is a function of the number of adults and environmental conditions. The asymptotic dynamic behaviour of the discrete population model is related to the steady-state solution of the continuous-time formulation. Conclusions include that there can be a lack of convergence to the steady-state age distribution in discrete event reproduction models. The iteroparous vital ratio (the ratio between the maximal age and the reproductive period) is fundamental to determining this convergence. When the vital ratio is rational, an equivalent discrete-time model for the population can be derived whose asymptotic dynamics are periodic and when there are a finite number of founder cohorts, the number of cohorts remains finite. When the ratio is an irrational number, effectively there is convergence to the steady-state age distribution. With a finite number of founder cohorts, the number of cohorts becomes countably infinite. The matrix model is useful to clarify numerical results for population models with continuous densities as well as delta measure age distribution. The applicability in ecotoxicology of the population matrix model formulation for iteroparous populations is discussed.     

Seasonal distribution of juvenile Atlantic cod in the southern Gulf of St Lawrence

Seasonal changes in the geographic distribution of juvenile (ages 1–4) Atlantic cod Gadus morhua were determined from research surveys conducted in the southern Gulf of St Lawrence during 1986–87, 1990–91, and 1994. From late June to early October, juvenile cod were most abundant in shallow water (<50 m deep). By early November, all age groups (adults and juveniles) had migrated to deep (>100 m) offshore waters for the winter. For most juveniles, this represented a minimum migration of >225 km (each way). Age 3 and 4 cod migrated greater distances than age 1 and 2 fish. Some age 3 and 4 cod migrated as far as 650 km from their summer and early autumn areas. Although there was partial geographic segregation of juvenile and adult fish during summer and winter, all age groups were found together during the migration. No cod were present in the shallow waters (<80 m deep) of the south-western Gulf during April due, in part, to later than average ice-melt. Large numbers of all age-classes were present in most of the south-eastern Gulf by mid-May.