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Global ocean ecosystem dynamics (GLOBEC) research has used a nested set of observations, experiments, and models in space and time to address the question of how climate change may affect marine populations. One of the great challenges has been to use observations effectively that span roughly 10 orders of magnitude spatially and temporally to understand variability in physical and biological environments. A major achievement has been the fostering of a coupled modelling and observational programme in a number of wellselected ecosystems globally. The advances in interdisciplinary observation and experimentation over the past decade, reviewed in this chapter, have led to signifi cant progress in understanding the structure and functioning of ocean ecosystems. The concept of target organisms (Gifford et al. , Chapter 4 , this volume) has been central to this approach and is linked to advances in the understanding of individual organism behaviours and population processes. New sampling and observation systems have been developed, particularly acoustic and optical. Shipboard, laboratory, and in situ process studies have been linked with new approaches to understanding trophic complexity. New approaches, applying techniques to retrospective studies, have contributed to understanding past ecosystem states and widespread use of a comparative approach has revealed new insights into the role of target species and the dynamics of marine ecosystems globally. [...]

We investigated the larval feeding ecology of sprat Sprattus sprattus L., an important commercial fish species in the Central Baltic Sea. We collected seasonally resolved larval abundance and gut content as well as zooplankton data, with the goal of identifying mechanisms leading to variability in larval survival. Our results show sprat larvae feed progressively on larger food items as they grow during the season depending on their size and the composition of the available prey. We found first-feeding larvae early in the spawning season to prey exclusively upon microplankton. Small larvae fed mainly upon nauplii of the copepods Acartia spp., Temora longicornis and Centropages hamatus (ca. 65%), whereas larger larvae consumed up to 80% Acartia spp. copepodites and adults, as well as cladocerans. Trends in sprat larval diets were to a large degree explainable by selective feeding. Feeding success and gut fullness increased linearly with larval size. Trophic niche breadth increased linearly until larvae reached a predator size of 16 mm, after which it decreased. We explain the latter decline by a restricted size spectrum of prey available to larger sprat larvae, which points towards the importance of considering the structure of the zooplankton community when evaluating the predator size to niche breadth relationship. Our results suggest first-feeding Baltic sprat larvae to be always food-limited, while larger larvae are not. We hypothesize medium-sized sprat larvae to be the life stage that has the potential to cause most of the interannual variability in sprat larval survival, which is dependent on a match between larval production and the state of the plankton cycle.

Large-scale climatic conditions prevailing over the central Baltic Sea resulted in declining salinity and oxygen concentrations in spawning areas of the eastern Baltic cod stock. These changes in hydrography reduced the reproductive success and, combined with high fishing pressure, caused a decline of the stock to the lowest level on record in the early 1990s. The present study aims at disentangling the interactions between reproductive effort and hydrographic forcing leading to variable recruitment. Based on identified key processes, stock dynamics is explained using updated environmental and life stage-specific abundance and production time-series. Declining salinities and oxygen concentrations caused high egg mortalities and indirectly increased egg predation by clupeid fish. Low recruitment, despite enhanced hydrographic conditions for egg survival in the mid-1990s, was due to food limitation for larvae, caused by the decline in the abundance of the copepod Pseudocalanus sp. The case of the eastern Baltic cod stock exemplifies the multitude effects climatic variability may have on a fish stock and underscores the importance of knowledge of these processes for understanding stock dynamics.

Oceanographic conditions in the brackish central Baltic Sea are strongly linked to atmospheric forcing and the unusual period of persistently strong westerlies that, since the late 1980s, have resulted in an increase in average water temperatures and decreasing salinity. These changes in temperature and salinity resulted in a change in the dominance of the mesozooplankton community from Pseudocalanus sp. to Temora longicornis and Acartia spp. Similar to the copepod community, the central Baltic fish community shifted from cod ( Gadus morhua ), dominant during the 1980s, to sprat ( Sprattus sprattus ), dominant during the 1990s. Further, the commercially important pelagic fish species herring ( Clupea harengus ) and sprat exhibited reductions in growth. Using Principal Component and Correlation Analyses we investigated the temporal variability in the importance of the food supply as well as competition on condition of central Baltic pelagic fish species. Our results indicate that herring condition results from a combined effect of changes in the food environment and increased competition with sprat, while sprat condition appeared to be primarily determined by intra-specific competition.

The recruitment processes of Baltic cod and sprat were analysed and critical periods were identified by addressing the major impact factors on individual early life history stages separately and relating observed abundance data between successive stages. For cod, recruitment appeared to be dependent on egg survival, with low oxygen concentration in dwelling depths and predation by clupeids as the major causes for egg mortality. Surviving egg production and larval abundance were weakly correlated, whereas larval abundance was significantly related to year class strength. This indicated that the period between the late egg and the early larval stage is critical for cod recruitment. A potential variable identified to affect this life stage was prey availability for larvae. For sprat, early and late egg stage production as well as late egg stage production and larval abundance were significantly related. However, year class strength was largely independent of larval abundance. Thus, the period between the late larval and early juvenile stage appeared to be critical for sprat recruitment. Potential variables identified to affect this life stage were ambient temperature and wind stress. Environmental factors showing statistically significant covariance with the survival of one of these critical life stages were incorporated into stock-recruitment models for individual spawning areas separately and for the Central Baltic combined.

Fish stock development in the Central Baltic Sea (1976-2000) in relation to variability in the environment - DTU Orbit (15/04/14) Fish stock development in the Central Baltic Sea (1976-2000) in relation to variability in the environment - DTU Orbit (15/04/14) Köster F, Möllmann C, Neuenfeldt S, Vinther M, St. John M, Tomkiewicz J et al. Fish stock development in the Central Baltic Sea (1976-2000) in relation to variability in the environment. I C E S Marine Science Symposia. 2003;219:294-306

Pseudocalanus elongatus is a key species in the pelagic zone of the deep basins of the Central Baltic Sea. The copepod serves as a major food organism for larval as well as for adult, pelagic planktivorous fish. Large interannual fluctuations in the standing stock of P. elongatus have been attributed to significant changes in the hydrographic environment over the last two decades. In particular, the decreasing salinity in the Baltic deep basins, a result of a change in atmospheric forcing leading to an increase in rainfall since the 1980s and of a lack of pulses of saline water intrusions from the North Sea, was found to affect reproduction and maturation of the copepod. In parallel, dramatic changes in the weight-at-age of herring, one of the most important commercial fishes of the Baltic Sea, have been observed since the late 1980s. Using time-series on herring stomach contents, as well as length and weight, we provide evidence for a chain of events relating variability in climate, salinity and P. elongatus abundance to changes in diet and condition of herring in the Central Baltic Sea.

Throughout the last 2 decades a shift from a cod- to a sprat-dominated system occurred in the upper trophic levels of the Central Baltic Sea. This was caused by a decline in the cod stock, due to recruitment failure and high fishing intensity, resulting in a decrease in predation pressure on sprat. Concurrently with the lowest cod stock size on record, sprat reached biomass values of above 2 × 106 t in 1992, being relatively stable afterwards. Besides predation mortality through cod and in recent years also an increasing fishing pressure, cannibalism on eggs may be a compensatory process limiting the reproductive success of sprat and hence contributing to the population regulation in the Central Baltic. Based on sprat stomach sampling on 21 cruises between March 1988 and July 1996 cannibalism on sprat eggs was investigated in the Bornholm Basin, one of the main spawning areas of Central Baltic sprat. Using a model of gastric evacuation to estimate daily food intake rates and a Virtual Population/Extended Survivor Analysis for computing predator population sizes, egg cannibalism rates were estimated. These were compared to egg abundance data from ichthyoplankton surveys and to preliminary estimates of seasonal egg productions. The study revealed significant interannual differences in the intensity of sprat egg cannibalism with considerable predation in 1990 to 1992 (>15% of the egg abundance during peak spawning and >60% of the seasonal production) and a reduction in most recent years (<16% of the corresponding abundance and production). As a possible reason for these differences a combination of changes in the vertical overlap of predator/prey and variability in the food environment were identified. Shortcomings of the applied methods and the possible impact of cannibalism on the reproductive success and population development of sprat in the Central Baltic Sea are discussed.

Long-term dynamics (1959–1997) of the copepod species Pseudocalanus elongatus, Temora longicornis, Acartia spp. and Centropages hamatus, as well as the taxonomic group of cladocerans, are described for the open sea areas of the central Baltic Sea. Differences between areas, i.e. Bornholm Basin, Gdansk Deep and Gotland Basin, as well as between 5 year periods, were investigated by means of Analysis of Variance (ANOVA). No significant differences in mesozooplankton biomass between areas were found. On the other hand, clear time-trends could be demonstrated and related to salinity and temperature, with P.elongatus biomass mainly dependent on salinity and T.longicornis, Acartia spp. and cladocerans biomasses dependent, to a large extent, on thermal conditions. Decreasing salinities since the early 1980s due to a lack of major inflows of highly saline water from the North Sea and increased river run-off, both triggered by meteorological conditions, obviously caused a decrease in biomass of P.elongatus. Contrarily, the standing stocks of the other abundant copepod species and cladocerans followed, to a large degree, the temperature development and showed, in general, an increase. The shift in species composition during this period is considered to be a reason for decreasing growth rates of Baltic herring (Clupea harengus) since the early 1980s, and for sprat (Sprattus sprattus) since the early 1990s. Generally, it is suggested that low mesozooplankton biomasses in the 1990s were caused, at least partially, by amplified predation by clupeid fish stocks.

Throughout the 1980s, reproductive success of the top-predator cod declined and stock sizes of the main prey species herring and especially sprat, important planktivorous predators in the system, increased substantially. Although the hydrographic conditions conducive for survival of early life stages improved during the 1990s, recruitment success of cod remained far below average. As clupeids have been identified as major predators on cod eggs and larvae in the Baltic, increased predation may be an important factor hampering stock recovery. Results from stomach content analysis of herring and sprat during the spawning season of 1988–1995 and ichthyoplankton surveys in the Bornholm Basin (the only important spawning area of cod in the Central Baltic in this period) allow a comparison of estimated consumption rates by the predator populations with standing stocks and production rates of cod eggs and larvae. Despite uncertainties in the estimation procedure, the findings confirm substantial predation on cod eggs by both clupeid species. Especially at the beginning of the cod-spawning season, characterized by low zooplankton availability, sprat consumed a considerable proportion of the eggs produced. In 1993, the relative importance of zooplankton as prey increased, while fish eggs were encountered more rarely. In contrast, predation by herring remained on the same level or even increased, especially late in the spawning season. The larval stage of cod is not substantially affected by predation owing to limited spatial overlap between prey and predator.

Considerable changes have taken place in the pelagic ecosystem of the Central Baltic Sea during the last decade. Owing to a combination of high fishing pressure and unfavourable hydrographic conditions, the cod (Gadus morhua) stock as the top predator in the system was reduced from a high level in the early 1980s to its lowest size on record in the early 1990s. The preferred prey species sprat (Sprattus sprattus) showed a significant increase in population size since the late 1980s to the highest level on record in recent years, while the herring (Clupea harengus), another important planktivore in the system, did not show such a response. We investigate whether fluctuations in clupeid stock size cascade down to the trophic level of mesozooplankton, based on stomach content data and daily ration estimates in combination with stock sizes estimated from Multispecies Virtual Population Analysis. Estimates of daily consumption by the populations of the two species for 1978–1990 were compared with standing stocks of mesozooplankton species. No evidence was found for food limitation as might be expected if clupeids exert a strong top-down control on mesozooplankton. Also no influence on interannual variability of mesozooplankton abundance was detected. However, predation did contribute to the seasonal development of two copepod species (Pseudocalanus elongatus and Temora longicornis).