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1.
The Baltic Sea ecosystem has undergone large changes during the last two decades, including a severe reduction in cod and herring biomass but, at the same time, a large increase in sprat abundance. The lower trophic levels of the Baltic Sea also changed due to environmental fluctuations, including variations in salinity and in volume of oxygenated water. In this apparently shifting environment, the conditions of herring and sprat have undergone large inter-annual variations during the past 15–20 years. In this study, we explore how abiotic factors (i.e. salinity and temperature) and biotic factors (biomass of the copepods Pseudocalanus elongatus , Temora longicornis , Acartia spp. and of cladocerans as well as clupeid abundance) in different seasons (May and August) affect clupeid body condition. Our analyses suggest that data of zooplankton biomass and abiotic factors in August have higher predictive power than May data. Although our analysis suggests that salinity (a bottom-up process) has an effect on sprat condition, total abundance of clupeids (a top-down process) is by far the most significant predictor of both herring and sprat condition. The strong correlation between clupeid abundance and total zooplankton biomass points to food competition and to top-down control by herring and sprat on common food resources. Furthermore, clupeid condition co-varied with the changes in the weight of zooplankton in the stomachs, which further suggest food competition being the main mechanism behind the changes in clupeid condition during the last two decades. Hence, our results are not in agreement with most of the current literature that has suggested that clupeid growth is regulated by environmentally mediated bottom-up processes acting on the abundance of copepods. This is, to our knowledge, the first evidence of food resources mediated density-dependent fish growth in a large marine ecosystem.  相似文献   

2.
The qualitative and quantitative composition of fish eggs and larvae in the stomach contents of Baltic herring and sprat was examined. The occurrence of fish eggs and larvae in the diet of herring was markedly higher, compared to sprat. Eggs of four and larvae of nine fish species were found in the herring stomachs. The prey species composition of the herring and sprat diets was related to the ichthyoplankton composition and to the degree of overlapping of the prey and predator distributions. Sprat eggs were the predominant prey in the herring diet in May and sprat larvae prevailed in August. The daily ration estimates indicate that the sprat larvae consumption by herring in August exceeded the sprat egg consumption in May by many times. Cod eggs in the herring stomachs were only found at the optimal oxygen content and salinity for the cod eggs in the near-bottom water layer.  相似文献   

3.
Predators commonly share prey with human exploiters, intuitively suggesting that there is an inherent human–predator conflict through competition for prey. Here we studied the effects of fishing and predation mortality on biomass distributions and yields of shared prey using a size‐structured model of competing populations, describing the life histories of Baltic Sea sprat and herring. Whereas both species responded in a similar fashion to increased fishing mortality, with decreasing juvenile and adult biomasses, we found that responses to predation mortality differed between species. Sprat only display weak compensatory responses with increasing predation mortality, while over a substantial range of mortalities there was a strong increase in adult (and total) herring biomass, i.e. overcompensation. The observed biomass overcompensation results from relaxed intraspecific competition as predation mortality increased, allowing for faster individual growth rates that in turn lead to a change in population composition (juvenile:adult biomass ratio). Our results suggest that the potential for biomass overcompensation is higher for species exhibiting substantial growth after maturation. Differences in size‐selectivity of predators and fishing mortality resulted in a positive effect of predation mortality on fisheries yields, which can be explained by an overcompensatory response in adult herring biomass. Thus, somewhat counter intuitive, our results suggest that fishermen, depending on prey life history, may actually benefit from allowing for a higher abundance of predators, despite competing for shared prey.  相似文献   

4.
Summary Multispecies virtual population analysis is an attempt to take species interactions into account when assessing the status of fish stocks. It was introduced primarily with the aim of lowering the uncertainty in the natural mortality rate as used in single species VPA and to take account of variability between years and ages by calculating inside the model the part of the mortality rate caused by predation. The output of the MSVPA is therefore —in addition to stock sizes and fishing mortality rates as in single species VPA —the amounts consumed of the various species by the predators included in the analysis.The MSVPA model of the predation interactions results in a set of coupled non-linear equations which must be solved for each time step in the analysis. Key parameters in the model are the so-called suitability coefficients, measuring the relative suitability of one species as prey for another species. These parameters must be estimated inside the model and this estimation requires data on the stomach contents of the predators in the model. The MSVPA makes two key assumptions: constant ration size (i.e. independent of time for each species-age combination) and hence fixed weights-at-age and a model of prey selection which leads to a type ii functional feeding response. These assumptions do not hold for all areas and therefore limit the applicability of the MSVPA in its present form.The MSVPA is probably one of the more successful multispecies models in fisheries. Its main application to date has been to the North Sea, and although it has not been used directly as a management tool it has provided input values of parameters used in assessment models as well as valuable insights into the system. For example, it has demonstrated that an increase in mesh size can result in lower long-term yields, an effect opposite to what is predicted if species interactions are ignored. Such insights into the dynamics of the system are useful and MSVPA may therefore have an indirect role to play in management. Nevertheless, due to many uncertainties involved in multispecies modelling in general and MSVPA in particular, it seems doubtful that the use of MSVPA in fisheries management will be much greater in the immediate future than it is at present.  相似文献   

5.
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.  相似文献   

6.
For the first time an international acoustic survey dataset covering three decades was used to investigate the factors shaping the spatial and temporal patterns in the condition of sprat and herring in the Baltic Proper. Generalized additive models showed that the spatial and temporal fluctuations in sprat density have been the main drivers of the spatio-temporal changes of both sprat and herring condition, evidencing intra- and inter-specific density dependence mediated by the size and distribution of the sprat population. Salinity was also an important predictor of herring condition, whereas temperature explained only a minor part of sprat model deviance. Herring density was an additional albeit weak significant predictor for herring condition, evidencing also intra-specific density dependence within the herring population. For both species, condition was high and similar in all areas of the Baltic Proper until the early 1990s, coincident with low sprat densities. Afterwards, a drop in condition occurred and a clear south–north pattern emerged. The drop in condition after the early 1990s was stronger in the northern areas, where sprat population increased the most. We suggest that the increase in sprat density in the northern areas, and the consequent spatial differentiation in clupeid condition, have been triggered by the almost total disappearance of the predator cod from the northern Baltic Proper. This study provides a step forward in understanding clupeid condition in the Baltic Sea, presenting evidence that density-dependent mechanisms also operate at the spatial scale within stock units. This stresses the importance of spatio-temporal considerations in the management of exploited fish.  相似文献   

7.
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.  相似文献   

8.
Population dynamics of major Baltic calanoid copepod speciesin the Gotland Basin during the last two decades were characterizedby a decline of Pseudocalanus elongatus associated with decliningsalinities, and an increase of Temora longicornis and Acartiaspp. potentially due to warmer conditions. Additionally thisstudy investigated the effect of predation by the major planktivorousfish species herring (Clupea harengus) and sprat (Sprattus sprattus)for the period 1977–1996 in the Gotland Basin (CentralBaltic Sea). Examination of consumption by these fish speciesin relation to copepod production estimates showed a switchby herring from consuming mainly CV/VI of P. elongatus and T.longicornis, to preying on CII of the latter copepod. This switchwas potentially due to increased competition with the drasticallyincreased sprat stock since the late 1980s. Further, an increasedpredation pressure by sprat on CV/CVI of both copepod speciesin spring resulted in higher copepod mortality rates. In consequence,based on these results we suggest that the increase in the spratstock since the late 1980s contributed to a decline of P. elongatus,and additionally prevented an even more pronounced temperature-drivenincrease in the T. longicornis stock, as was observed for Acartiaspp., which was not significantly consumed.  相似文献   

9.
In 2014, the International Council for the Exploration of the Sea (ICES) was unexpectedly unable to provide an analytical assessment of eastern Baltic cod stock; factors such as data issues, assessment methodology, and the ecological situation of cod were indicated as the reasons for this failure. Some evidence suggests that the natural mortality (M) of cod could increase substantially in forthcoming years and that the selectivity could change. In this paper, age‐structured and stock‐production assessment models were applied to simulate the dynamics of cod stock; in the models, both constant and increasing natural mortalities were permitted. In the age‐structured model, the effects of selectivity related to the cod size on the cod assessment were also analysed. In addition, stock with characteristics similar to Baltic cod stock and increasing natural mortality was generated and assessed with the age‐structured model using both constant and increasing M. It was shown that models with increasing natural mortality of cod in recent years perform much better than models with constant natural mortality in terms of the distribution of residuals and retrospective patterns. The models with size‐dependent selectivity did not perform better than other standard assessments. The assessment of generated stock (where natural mortality was increasing) with constant natural mortality in the assessment model showed a poor distribution of residuals and strong retrospective patterns, similar to the ICES assessment with constant M. The conducted simulations strongly suggest that the main reason for the poor recent cod assessment is the increase in natural mortality, which is not considered in the assessment methodology.  相似文献   

10.
This paper updates previous reviews of the 1993 stock decline of Pacific herring (Clupea pallasi) in Prince William Sound, Alaska, and focuses on hypotheses about subsequent poor recovery. Recent age structured assessment modeling with covariate analysis indicates that the population dynamics of the sound’s herring are influenced by oceanic factors, nutrition, and, most substantially, hatchery releases of juvenile pink salmon. For the 1993 decline, poor nutrition remains the most probable cause with disease a secondary response. Concerning poor recovery, we examined 16 potential factors and found three to be causal: oceanic factors, poor nutrition, and hatchery releases of juvenile pink salmon. Absences of strong year classes at both Sitka and Prince William Sound after 1993 indicate the action of large-scale ocean processes. Beyond regional-scale environmental factors, two factors specific to the sound influence the population dynamics of herring and are likely impeding recovery. First, pink salmon fry releases have increased to about 600 million annually and may disrupt feeding in young herring, which require adequate nutrition for growth and overwintering survival. Juvenile pink salmon and age-1 herring co-occur in nearshore areas of bays in late spring and summer, and available data on dietary overlap indicates potential competition between the age-1 juvenile herring and juvenile pink salmon. Field studies demonstrate that juvenile herring reduce food intake substantially in the presence of juvenile pink salmon. Second, overwintering humpback whales may consume potentially large amounts of adult herring, but further studies must confirm to what extent whale predation reduces herring biomass.  相似文献   

11.
Feeding ecology of central Baltic Sea herring and sprat   总被引:7,自引:0,他引:7  
A unique dataset of stomach contents sampled between 1977 and 1999 in the central Baltic Sea was used to perform a comprehensive study of the feeding ecology of Central Baltic herring Clupea harengus and sprat Sprattus sprattus . Both fish species were mainly preying upon calanoid copepods with Pseudocalanus sp. dominating the diet of herring, whereas sprat generally preferred Temora longicornis . Sprat preyed upon older copepodite stages, indicating size‐selective particulate feeding, whereas herring additionally fed on smaller copepodite stages, indicating occasional low food supply inducing filter‐feeding. Additional food sources other than copepods were mysids in winter and autumn for medium to large herring, as well as cladocerans for sprat in spring and summer, determined by the seasonal occurrence of these plankton species. Seasonally the highest feeding activity of both fishes species occurred in spring and summer, the main reproductive periods of calanoid copepods. The most important food item for both predators in spring was Pseudocalanus sp. In summer sprat switched to T. longicornis and Acartia spp. Since the late 1970s, the total stomach fullness decreased and the fraction of empty stomachs increased. In parallel the amount of Pseudocalanus sp. in the diets of both fish species decreased. Further, a considerable dietary overlap between both species in spring indicated considerable competition for food resources, especially due to an enlarged sprat stock. The results of this study support the hypothesis that growth reductions observed in Baltic herring and sprat are due to combination of a change in food availability and an increase in density‐dependent competition.  相似文献   

12.
The consumption of estuarine copepods by juvenile herring and sprat during estuarine residency was estimated using fish biomass data and daily rations calculated from two models of feeding in fish: a bioenergetic model and a gastric evacuation model. The bioenergetic model predicted daily rations that were, on average, three times higher than those estimated by a model based on field records of stomach contents. The biomass of herring and sprat in the estuary was negatively correlated with the daily ration suggesting that the clupeid fish populations were resource-limited. Copepod production decreased towards the winter and peaked in spring and summer. The relative importance of predation changed seasonally in function of the migration pattern of herring and sprat. In the spring and the summer, in situ production ofcopepod biomass was higher than the in situ consumption by fish. During the fall and the winter, consumption exceeded production. This suggests that top–down control exerted by marine pelagic fish may be an important force structuring estuarine copepod populations.  相似文献   

13.
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.  相似文献   

14.
The Baltic Sea is characterised by a heterogeneous oceanographic environment. The deep water layers forming the habitat of Baltic cod (Gadus morhua callarias L.) are subjected to frequently occurring pronounced anoxic conditions. Adverse oxygen conditions result in physiological stress for organisms living under these conditions. For cod e.g. a direct relationship between oxygen availability and food intake with a decreasing ingestion rate at hypoxia could be revealed. In the present study, the effects of oxygen deficiency on consumption rates were investigated and how these translate to stock size estimates in multi‐species models. Based on results from laboratory experiments, a model was fitted to evacuation rates at different oxygen levels and integrated into the existing consumption model for Baltic cod. Individual mean oxygen corrected consumption rates were 0.1–10.9% lower than the uncorrected ones. At the currently low predator stock size, however, the effect of oxygen‐reduced consumption on the total amount of eaten prey biomass and thus predation mortalities was only marginal. But should successful management lead to higher cod stock sizes in the future, then total predation mortalities will greatly increase and thus improved precision of these estimates would be valuable for the assessment of prey stocks.  相似文献   

15.
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.  相似文献   

16.
Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.  相似文献   

17.
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.  相似文献   

18.
Climate change and overfishing may lead to ecosystem instability and may benefit nonexploited organisms such as jellyfish. In the Irish Sea, an increase in jellyfish abundance was evident (r2=0.29, P=0.03) in a 16‐year time‐series (1994–2009) collected during juvenile fish surveys. Jellyfish abundance correlated positively with sea surface temperature (SST) over the preceding 18 months (r=0.65, pACF<0.001) and copepod biomass in the previous year (r=0.56, pACF=0.03) and negatively with spring (February–May) precipitation (r=?0.57, pACF=0.02). Principal components regression indicated that climatic indices explained 68% of the interannual variability in jellyfish abundance (P=0.003), where the components were based on the North Atlantic Oscillation Index, SST and precipitation. The frequency of cnidarian material present in Continuous Plankton Recorder (CPR) samples has also increased since 1970, with a period of frequent outbreaks between 1982 and 1991. Before this period, the herring stock in the northern Irish Sea declined rapidly to a low level, potentially stimulating structural change in the ecosystem. In 1985, there was a step decrease in CPR copepod biomass and in 1989, a step increase in the phytoplankton colour index, suggesting a cascading regime shift during the 1980s. Subsequent overexploitation of gadids, coupled with warm temperatures and the poor recruitment of cod, led to the rapid decline in cod biomass from 1990. While the biomass of sprat has decreased in the last decade, the herring stock has recovered partially. Reductions in demersal fishing pressure since 2000, intended to stimulate cod recovery, appear to have facilitated further rises in haddock biomass. Since the 1980s regime shift, sea temperatures have increased, the fish community has altered and jellyfish abundance has risen such that jellyfish and haddock may now play an increasingly important role in the ecosystem.  相似文献   

19.
20.
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.  相似文献   

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