Feeding ecology of pelagic fish species in the Gulf of Riga (Baltic Sea): the importance of changes in the zooplankton community

The feeding ecology of four pelagic fish species was studied in relation to their prey availability in the Gulf of Riga (Baltic Sea) during the summer 1999-2006. The zooplankton community was dominated by the cladoceran Bosmina longispina, rotifers Keratella cochlearis and K. quadrata and the copepod Eurytemora affinis, with the highest interannual variability in abundance recorded for B. longispina. The last influenced the diet of adult sprat Sprattus sprattus, juvenile smelt Osmerus eperlanus and three-spined stickleback Gasterosteus aculeatus as these were strongly selecting for B. longispina. The fish feeding activity did not match the abundance dynamics of their preferred prey, suggesting that fishes may switch to consume other prey in case the preferred diet was limited. A considerable dietary overlap indicated high potential competition between pelagic fish species. While herring Clupea harengus membras and G. aculeatus were relying on very different food, the diets of young O. eperlanus and G. aculeatus were very similar. Interannual variability in zooplankton composition and abundance significantly affected the diet composition of fishes, but those changes were insufficient to exert a consistent influence upon fish feeding activity and total amounts of zooplankton consumed.     

Inter‐annual variation in herring,Clupea harengus,and sprat,Sprattus sprattus,condition in the central Baltic Sea: what gives the tune?

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.     

Wasp-waist interactions in the North Sea ecosystem

Background

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Methodology/Principal Findings

We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill.

Conclusion/Significance

Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that “wasp-waist” interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.     

Stomach analyses of Baltic salmon from 1959–1962 and 1994–1997: possible relations between diet and yolk-sac-fry mortality (M74)

In recent years, Baltic Sea salmon Salmo salar have suffered high larval mortality (M74) which can be cured by thiamine treatment. Analyses of long term mortality records (1928–1998) from two salmon hatcheries suggest that before the 1970s M74 did not occur, or was less frequent. This indicates that varying M74 did not cause the long-term fluctuations of Baltic salmon catches in this period. The frequency of M74 has been correlated positively to the abundance of the salmon's primary prey, sprat Sprattus sprattus. Sprat, herring Clupea harengus and three-spined stickleback Gasterosteus aculeatus dominated the diet in both the periods of study, 1959–1962 and 1994–1997. The mean size of consumed sprat was significantly smaller in 1994–1997 compared with 1959–1962. Herring and, to a lesser extent, three-spined stickleback increased in the diet of salmon, while sprat appeared to constitute a smaller part of the diet in 1994–1997. The cause of M74 and the thiamine deficiency involved remains unknown, but is thought to be related to changes in thiamine or thiaminase content in forage fish, winter-feeding of salmon or general changes in the pelagic food web, caused by overfishing or eutrophication.     

Factors controlling long-term changes of the eutrophicated ecosystem of Pärnu Bay,Gulf of Riga

Phytoplankton, mesozooplankton, mysids and fish larvae were studied during 15–29 annual cycles measured weekly to monthly in Pärnu Bay, the Gulf of Riga. The monthly variability of the biological data was related to temperature, ice conditions, salinity, influx of nutrients, the North Atlantic Oscillation (NAO) index, cloudiness and solar activity. Phytoplankton development was mainly a function of the NAO index. For the whole study period the abundance of zooplankton increased with increasing water temperature and solar activity. Significant correlations between phytoplankton and zooplankton densities were found until 1990. After the invasion of the predatory cladoceran Cercopagis pengoi in 1991, the zooplankton community was likely to be regulated by the introduced species rather than phytoplankton dynamics. The increased abundances of rotifers and copepods triggered the increase in mysid densities. The development of herring larvae was positively affected by the high density of copepods and rotifers but also by increased eutrophication. Until 1990 there was no significant relationship between the density of zooplankton and herring larvae. A negative relationship between the density of zooplankton and herring larvae in the 1990s suggests that the major shift in zooplankton community resulted in food limitation for herring larvae. The results indicated that (1) atmospheric processes in the northern Atlantic explain a large part of the interannual variation of the local phytoplankton stock, (2) trophic interactions control the development of pelagic communities at higher trophic levels, and (3) the introduction of an effective intermediate predator has repercussions for the whole pelagic food web in Pärnu Bay.     

The effect of prey quality and ice conditions on the nutritional status of Baltic gray seals of different age groups

We analyzed a long-term data set of the body condition of Baltic gray seals (Halichoerus grypus) over time and investigated how average subcutaneous blubber thickness of different age groups of seals corresponds to environmental factors. Blubber thickness of pups declined until 2010. The decreasing weight of 5–6-year-old herring (Clupea harengus), the main prey fish for Baltic gray seals, explained well the decline. In the Gulf of Finland, the blubber thickness of pups declined also in recent years (2011–2015) with declining number of days with permanent ice cover. In other regions, the blubber thickness of pups increased during recent years with increasing weight of herring. The blubber thickness of sub-adults in Baltic Proper and that of hunted adult females in the Bothnian Bay also increased during recent years, and the weight of age 6+ or 7-year-old herring best explained the increase. The blubber thickness of all age groups of seals was thinnest in the Bothnian Bay where also herring weight was lowest. There was a negative correlation between blubber thickness of seals and herring catch size (an index of herring abundance) suggesting that herring quality, not the quantity, is important for the nutritional status of Baltic gray seals. Nutritional status of gray seals may thus reveal changes in the marine food web which affect herring quality. Marine food web, in turn, may be affected, e.g., by climate change. The warming climate also has an impact on ice cover and thus body condition of seal pups.     

Larval fish body growth responses to simultaneous browning and warming

Organisms are facing global climate change and other anthropogenic pressures, but most research on responses to such changes only considers effects of single drivers. Observational studies and physiological experiments suggest temperature increases will lead to faster growth of small fish. Whether this effect of warming holds in more natural food web settings with concurrent changes in other drivers, such as darkening water color (“browning”) is, however, unknown. Here, we set up a pelagic mesocosm experiment with large bags in the Baltic Sea archipelago, inoculated with larval Eurasian perch (Perca fluviatilis) and zooplankton prey and varying in temperature and color, to answer the question how simultaneous warming and browning of coastal food webs impact body growth and survival of larval perch. We found that browning decreased body growth and survival of larval perch, whereas warming increased body growth but had no effect on survival. Based on daily fish body growth estimates based on otolith microstructure analysis, and size composition and abundance of available prey, we explain how these results may come about through a combination of physiological responses to warming and lower foraging efficiency in brown waters. We conclude that larval fish responses to climate change thus may depend on the relative rate and extent of both warming and browning, as they may even cancel each other out.     

Tendencies in the Herring Population Development of the Baltic Sea

During the 1960's a change in population structure of Baltic herring started which resulted in the dominance of spring spawners in the entire Baltic since the beginning 1970's. Autumn spawning herring is very rare in the yields of fisheries since. This development has been accompanied by a likely increase of the total stock biomass of herring in the area. Yields of herring fisheries increased remarkable up to 1984, partly as a result of increased fishing effort. Recent developments of stock biomasses point to dependencies on fluctuations of growth rates. Growth is influenced by several environmental factors but is very likely especially dependent on abundance of food and on temperature. Eutrophication of the Baltic Sea increased perhaps the abundance of food for the planktonfeeding herring but it may have been contributing to the depletion of autumn spawning herring via the declining oxygen content of bottom water layers during the past 20 years.     

Trophic interactions involving mysid shrimps (Mysidacea) in the near-bottom habitat in the Baltic Sea

In a six month mesocosm tank experiment, hypotheses were tested concerning the role of benthopelagic mysid shrimps (Mysidacea) in the near-bottom food web of the Bothnian Sea, in the northern Baltic Sea. The first hypothesis tested was that the mysids interact, through predation, with benthic deposit-feeding Monoporeia affinis amphipods. A second hypothesis tested was that the sediment type is important for the overwintering success of the mysids. Changes in abundance and mass were recorded for M. affinis and mysids when separate and when coexisting, in two sediment types differing in organic content (food level); soft muddy clay (rich) and fine sand (poor). Despite the fact that newborn M. affinis offspring, a plausible target for predation by mysids, were present in substantial numbers in the tanks, no consistent evidence for any interaction between these taxa was found. The biomass of mysids was slightly higher in the muddy clay than in the sand tanks, and the mechanism behind this substrate effect is discussed. A third hypothesis, that the mysids interact with near-bottom zooplankton, was investigated. The tanks were continually supplied with in situ near-bottom sea-water containing a semi-natural assemblage of near-bottom plankton. As a result of mysid predation, tanks with mysids had lower abundance and biomass of cyclopoid copepods than tanks without mysids. Thus, the major interaction found was predation on near-bottom zooplankton by mysids and it is suggested that this interaction could potentially be an important food link, especially during periods with low food availability in the pelagic system.     

Impacts of elevated terrestrial nutrient loads and temperature on pelagic food‐web efficiency and fish production

Both temperature and terrestrial organic matter have strong impacts on aquatic food‐web dynamics and production. Temperature affects vital rates of all organisms, and terrestrial organic matter can act both as an energy source for lower trophic levels, while simultaneously reducing light availability for autotrophic production. As climate change predictions for the Baltic Sea and elsewhere suggest increases in both terrestrial matter runoff and increases in temperature, we studied the effects on pelagic food‐web dynamics and food‐web efficiency in a plausible future scenario with respect to these abiotic variables in a large‐scale mesocosm experiment. Total basal (phytoplankton plus bacterial) production was slightly reduced when only increasing temperatures, but was otherwise similar across all other treatments. Separate increases in nutrient loads and temperature decreased the ratio of autotrophic:heterotrophic production, but the combined treatment of elevated temperature and terrestrial nutrient loads increased both fish production and food‐web efficiency. CDOM: Chl a ratios strongly indicated that terrestrial and not autotrophic carbon was the main energy source in these food webs and our results also showed that zooplankton biomass was positively correlated with increased bacterial production. Concomitantly, biomass of the dominant calanoid copepod Acartia sp. increased as an effect of increased temperature. As the combined effects of increased temperature and terrestrial organic nutrient loads were required to increase zooplankton abundance and fish production, conclusions about effects of climate change on food‐web dynamics and fish production must be based on realistic combinations of several abiotic factors. Moreover, our results question established notions on the net inefficiency of heterotrophic carbon transfer to the top of the food web.     

Stable isotopes show food web changes after invasion by the predatory cladoceran <Emphasis Type="Italic">Cercopagis pengoi</Emphasis> in a Baltic Sea bay

Cercopagis pengoi, a recent invader to the Baltic Sea and the Laurentian Great Lakes, is a potential competitor with fish for zooplankton prey. We used stable C and N isotope ratios to elucidate trophic relationships between C. pengoi, zooplankton (microzooplankton, 90–200 m, mostly copepod nauplii and rotifers; mesozooplankton, >200 m, mostly copepods), and zooplanktivorous fish (herring, size range 5–15 cm and sprat, 9–11 cm) in a coastal area of the northern Baltic Sea. The isotope ratios in C. pengoi and fish were much higher than those of zooplankton, showing general trends of enrichment with trophic level. Young-of-the-year (YOY) herring had a significantly higher ¹⁵N/¹⁴N ratio than C. pengoi, suggesting of a trophic linkage between the two species. To evaluate the possible relative importance of different food sources for C. pengoi and YOY herring, two-source isotope-mixing models for N were used, with micro- and mesozooplankton as prey for C. pengoi and mesozooplankton and C. pengoi as prey for YOY herring. These models indicate that mesozooplankton was the major food source of both species. However, microzooplankton may be important prey for young stages of C. pengoi. Comparative analyses of the herring trophic position before and after the invasion by C. pengoi showed a trophic level shift from 2.6 to 3.4, indicating substantial alterations in the food web structure. Our findings contribute to a growing body of evidence, showing that C. pengoi can modify food webs and trophic interactions in invaded ecosystems.     

Spatially structured interactions between a migratory pelagic predator, the Norwegian spring-spawning herring Clupea harengus L., and its zooplankton prey

Spring-spawning herring Clupea harengus was patchily distributed over large parts of the Norwegian Sea in May 1995–2005, during the early phase of the annual feeding migration. Overall, herring tended to be found in areas with intermediate biomasses of zooplankton prey, intermediate water temperatures and relatively high salinities. Herring had more food in their stomachs in areas of relatively low water temperature and high herring abundance. Hydrographical conditions revealed that herring was feeding mainly within Atlantic water masses, and more intensely in western and northern regions of the Norwegian Sea. Zooplankton biomass was patchily distributed, and was generally higher towards the western parts of the Norwegian Sea. Here, zooplankton biomass in year _{i +1} was also negatively associated with herring spawning stock biomass in year _i , while there was no evidence for such an association in the eastern region; indicating that herring may have a geographically structured 'top–down' effect on the recruitment of its zooplankton prey. The fact that herring was not typically associated with the areas containing the greatest zooplankton biomasses may reflect that the fish had not yet reached the most profitable feeding grounds or alternatively that herring was depleting zooplankton biomass.     

Multi-level trophic cascades in a heavily exploited open marine ecosystem

Anthropogenic disturbances intertwined with climatic changes can have a large impact on the upper trophic levels of marine ecosystems, which may cascade down the food web. So far it has been difficult to demonstrate multi-level trophic cascades in pelagic marine environments. Using field data collected during a 33-year period, we show for the first time a four-level community-wide trophic cascade in the open Baltic Sea. The dramatic reduction of the cod (Gadus morhua) population directly affected its main prey, the zooplanktivorous sprat (Sprattus sprattus), and indirectly the summer biomass of zooplankton and phytoplankton (top-down processes). Bottom-up processes and climate-hydrological forces had a weaker influence on sprat and zooplankton, whereas phytoplankton variation was explained solely by top-down mechanisms. Our results suggest that in order to dampen the occasionally harmful algal blooms of the Baltic, effort should be addressed not only to control anthropogenic nutrient inputs but also to preserve structure and functioning of higher trophic levels.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Food-web indicators accounting for species interactions respond to multiple pressures

Food-web indicators for marine management are required to describe the functioning and structure of marine food-webs. In Europe, the Marine Strategy Framework Directive (MSFD), intended to lead to a ‘good environmental status’ of the marine waters, requires indicators of the status of the marine environment that also respond to manageable anthropogenic pressures. Identifying such relationships to pressures is particularly challenging for food-web indicators, as they need to be disentangled from linkages between indicators of different functional groups caused by species interactions. Still, such linkages have not been handled in the indicator development. Here we used multivariate autoregressive time series models to identify how fish indicators in an exploited food-web relate to fishing, climate and eutrophication, while accounting for the linkages between indicators caused by species interactions. We assembled 31-year long time series of indicators of key functional groups of fish in the Central Baltic Sea pelagic food-web, which is characterized by strong trophic links between cod (Gadus morhua) and its main fish prey sprat (Sprattus sprattus) and herring (Clupea harengus). These food-web indicators were either abundance-based indicators of key piscivores (cod) and zooplanktivores (sprat and herring) or size-based indicators of the corresponding trophic groups (biomass of large predatory fish (cod ≥ 38 cm) and biomass of small prey fish (sprat and herring <10 cm)). Comparative analyses of models with and without linkages among indicators showed that for both types of indicators, linkages corresponding to predator-prey feedbacks and intra-specific density-dependence were essential to explain temporal variation in the indicators. Thus, no indicator-pressure relationships could be found that explained the indicators’ variation unless such linkages were accounted for. When accounting for these, we found that the indicators overall respond to multiple pressures acting simultaneously rather than to single pressures, as no pressure alone could explain how the indicators developed over time. The manageable pressures fishing and eutrophication, as well as the prevailing hydrological conditions influenced by climate, were all needed to reproduce the inter-annual changes in these food-web indicators combined, although individual relationships differed between the indicators. We conclude that our innovative indicator-testing framework can therefore be used to identify responses of food-web indicators to manageable pressures while accounting for the biotic interactions in food-webs linking such indicators.     

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of ¹³C by the copepods as a proxy for carbon uptake in feeding experiment with ¹³C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.     

Towards better Understanding of Successful Management of the Gulf of Riga Herring Stock

The Baltic herring is one of the key elements of the pelagic ecosystem of the Baltic Sea, being of the most important commercial species of the sea. The structure and dynamics of herring populations (stocks) have been assessed and managed on international level since 1970s. Since 1990, all local herring populations have been combined and assessed as one stock (Central Baltic Herring in Sub-divisions 25–29 and 32). However, the continuously decreasing trend in stock biomass throughout almost 30-years period of observations indicates the failure of the stock management implementation of the scientific advice. The separate assessments of different stocks and results of hydro-acoustic surveys of different sub-units (herring in Sub-divisions 25–27, Sub-divisions 28, 29 and 32 and the Gulf of Riga herring) have revealed rather different situation within combined assessment unit: e.g. fishing mortality of the Gulf of Riga herring has decreased while a sharp increase in mortality is observed in the herring stocks of the North-eastern Baltic (Sub-divisions 28, 29 and 32). At the same time stock abundance and biomass of the Gulf of Riga herring have increased to record high in early 2000s allowing also higher catches. It is concluded that only separate analytical assessment of local populations combined with regular acoustical surveys and following different protection measures have proved to be a successful way in managing the herring stocks. Joint assessment and management of several populations with different structure and dynamics as one combined stock do not allow revealing the real situation and trends in its parts.     

Long-term dynamics of zooplankton in the southeastern Baltic Sea

Long-term research in the Baltic Sea revealed the basic trends of zooplankton community variations depending on oceanographic processes. Alternation of the periods of increase and decrease in salinity of the Baltic Sea against the background of climate changes (temperature increase) and eutrophication affect the state of the entire Baltic ecosystem, including zooplankton. For these periods, the dynamics of zooplankton in the Baltic Sea were analyzed based on literature data and results of regular research in the southeastern Baltic Sea during 1998–2007. The changes in the hydrological situation were accompanied by significant changes in the zooplankton community. In the 1990s–2000s, the abundance and biomass of brackish-water and thermophilous species primarily of Cladocera and Copepoda increased markedly. The role of the previously dominant marine copepod Pseudocalanus elongatus decreased due to salinity reduction in the deep-water part of the Baltic Sea. Maximum development of zooplankton occurred in years of the greatest warming-up of the water (2001, 2005–2007) against the background of a general positive trend of zooplankton abundance in the last decade.