Different growth trends of whitefish (Coregonus lavaretus) forms in the northern Baltic Sea

The sea growth of two whitefish forms, anadromous (Coregonus lavaretus lavaretus) and sea‐spawning (Coregonus lavaretus widegreni), was analysed using samples collected from the commercial sea catch in the Gulf of Bothnia (GoB) in the northern Baltic Sea during 1998–2014. In the GoB area, these two forms are possible to identify because the gill‐raker number and size at maturity vary between forms. The growth rate of the forms is linked to their feeding area. Sea‐spawning whitefish, which has a feeding migration near its home site, was shorter in the northern GoB (66°N–64°N) at the ages of 3–11 than those in the southern GoB (64°N–60°30′N). In the data, most whitefish were caught with gill nets in the GoB. The mesh sizes of gill nets capturing the anadromous form were mostly 35–45 mm, while those capturing the sea‐spawning form were <35 mm in the northern GoB. It is likely that the different growth trends for small and large whitefish were connected with differences in their recruitment for fishing. The length of anadromous females at the age of four sea years increased significantly, but the length of six‐year‐old anadromous female whitefish decreased over the catch years from 1998–2014. In contrast, the length of slow‐growing sea‐spawning whitefish of six years or older increased significantly in relation to the catch year in the gill‐net catch. The increase in the growth of young age groups in both forms was probably associated with the increasing temperature and the low fishing pressure on small fish. The decreasing age at capture for both forms and the depression of the mean size of old anadromous whitefish are signs of high fishing pressure with a high gill‐net effort that selectively removes the largest and oldest individuals of both forms.     

Generation of a neutral FST baseline for testing local adaptation on gill raker number within and between European whitefish ecotypes in the Baltic Sea basin

Divergent selection at ecologically important traits is thought to be a major factor driving phenotypic differentiation between populations. To elucidate the role of different evolutionary processes shaping the variation in gill raker number of European whitefish (Coregonus lavaretus sensu lato) in the Baltic Sea basin, we assessed the relationships between genetic and phenotypic variation among and within three whitefish ecotypes (sea spawners, river spawners and lake spawners). To generate expected neutral distribution of F_ST and to evaluate whether highly variable microsatellite loci resulted in deflated F_ST estimates compared to less variable markers, we performed population genetic simulations under finite island and hierarchical island models. The genetic divergence observed among (F_CT = 0.010) and within (F_ST = 0.014–0.041) ecotypes was rather low. The divergence in gill raker number, however, was substantially higher between sea and river spawners compared to observed microsatellite data and simulated neutral baseline (P_CT > F_CT). This suggests that the differences in gill raker number between sea and river spawners are likely driven by divergent natural selection. We also found strong support for divergent selection on gill raker number among different populations of sea spawners (P_ST > F_ST), most likely caused by highly variable habitat use and diverse diet. The putative role of divergent selection within lake spawners initially inferred from empirical microsatellite data was not supported by simulated F_ST distributions. This work provides a first formal test of divergent selection on gill raker number in Baltic whitefish, and demonstrates the usefulness of population genetic simulations to generate informative neutral baselines for P_ST–F_ST analyses helping to disentangle the effects of stochastic evolutionary processes from natural selection.     

Impact of 21st century climate change on the Baltic Sea fish community and fisheries

The Baltic Sea is a large brackish semienclosed sea whose species-poor fish community supports important commercial and recreational fisheries. Both the fish species and the fisheries are strongly affected by climate variations. These climatic effects and the underlying mechanisms are briefly reviewed. We then use recent regional – scale climate – ocean modelling results to consider how climate change during this century will affect the fish community of the Baltic and fisheries management. Expected climate changes in northern Europe will likely affect both the temperature and salinity of the Baltic, causing it to become warmer and fresher. As an estuarine ecosystem with large horizontal and vertical salinity gradients, biodiversity will be particularly sensitive to changes in salinity which can be expected as a consequence of altered precipitation patterns. Marine-tolerant species will be disadvantaged and their distributions will partially contract from the Baltic Sea; habitats of freshwater species will likely expand. Although some new species can be expected to immigrate because of an expected increase in sea temperature, only a few of these species will be able to successfully colonize the Baltic because of its low salinity. Fishing fleets which presently target marine species (e.g. cod, herring, sprat, plaice, sole) in the Baltic will likely have to relocate to more marine areas or switch to other species which tolerate decreasing salinities. Fishery management thresholds that trigger reductions in fishing quotas or fishery closures to conserve local populations (e.g. cod, salmon) will have to be reassessed as the ecological basis on which existing thresholds have been established changes, and new thresholds will have to be developed for immigrant species. The Baltic situation illustrates some of the uncertainties and complexities associated with forecasting how fish populations, communities and industries dependent on an estuarine ecosystem might respond to future climate change.     

Selectivity of fishing gears used in the Baltic Sea cod fishery

The Baltic cod (Gadus morhua) plays a very significant role in the Baltic Sea ecosystem being the major fish top predator and the most important commercial species for the Baltic Sea fishing industry. Consequently the management and understanding of the exploitation pattern of the stock is of major importance. Technical regulations, improving the selectivity of cod, have been a major management strategy and the Baltic Sea is likely to be the area where most fishing gear selectivity studies, focussing on size selectivity, have been conducted over time. The methodology for conducting and analysing selectivity data has been significantly improved in recent years. The subject is reviewed since the choice of methodology can have a significant effect on the interpretation of the outcome of selectivity experiments. Factors affecting the selectivity of trawl and gill nets are considered. Alternative ways to improve the size and species selectivity of trawls using selective devices are reviewed. Selectivity parameters from available literature are listed and the correlations of selectivity parameters to the mesh size for different gears are estimated. The historical legislation on selectivity is reviewed and the expected selectivity for trawls is estimated. Management considerations concerning the mortality of escaping and discarded fish and wider management impacts have to be considered if improving selectivity. The review is ended by conclusions including reflections on needed research in the future.     

Trace fossils as indicators of environmental change in Holocene sediments of the Archipelago Sea, northern Baltic Sea

Biogenic structures in Holocene sediments from the Archipelago Sea, northern Baltic Sea, were characterized through the analysis of X-ray images of four cores collected from water depths between 32 and 66 m. In the area, initial colonization of endobenthic invertebrates occurred at 7800 ± 80 calendar years BP. The trace assemblage at this level is low-diversity, small-diameter, shallowly tiered, and Palaeophycus-dominated; rare Arenicolites are also observed. Early colonization coincides with increasing marine influence in the post-glacial lacustrine setting just before the dramatic onset of brackish-water conditions established after 7600 BP. The post-incursion brackish-water assemblage possesses a higher diversity of traces (Planolites, Arenicolites, Lockeia, Teichichnus), which is taken to reflect the enhanced salinity and trophic state of the basin. The shift from Palaeophycus-mottling to a Planolites-dominated fabric represents changed behavioural patterns in the endobenthic community due to changed substrate properties. The ethology of the succeeding trace assemblage also represents a switch from domicile-based activities, such as predation, scavenging and interface-dominated deposit feeding to shallow-tier deposit feeding. Finally, traces are excluded from thinly laminated intervals, demonstrating that seafloor oxygen deficiency commonly reached levels that were detrimental to colonization of the sediment substrate.     

First observation of microcystin-LR in pelagic cyanobacterial blooms in the northern Baltic Sea

Cyanobacterial bloom samples from the Gulf of Finland (northern Baltic Sea) were collected in July 2003 and analyzed for microcystins and nodularins, cyanobacterial peptide hepatotoxins, by ELISA, HPLC-UV and LC-MS. The blooms consisted mainly of the genera Nodularia, Anabaena and Aphanizomenon. The main hepatotoxin in the samples was nodularin-R (Nod-R), all the samples also contained demethylnodularin-R. The presence of microcystin-LR was confirmed in three locations out of nine by multiple reactant monitoring on the triple quadrupole mass spectrometer. This is the first reported finding of microcystins in the Baltic Sea from the open sea area. Anabaena was the likely producer of microcystin-LR in the samples.     

Annual variation in gobiid larval density in the northern Baltic Sea

The highest densities of gobiid larvae along the Finnish Baltic coast from 1974 to 1996 were found in the Åland Islands and the Archipelago Sea. The annual variation in gobiid densities was three-fold (from 1· 1 to 3·3 larvae m⁻²) with peak densities in the years 1977 and 1992 and the lowest density in 1987. No long-term trends in the density were found. The earliest gobiids were caught in the first half of May and the occurrence of larvae peaked at the end of June to early July.Within breeding seasons there was a positive correlation between water temperature at a depth of 1 m and the density of larvae. The density of larvae, however, did not correlate with mean summer temperature or the mean temperature of the previous summer. Instead larvae production was negatively affected by the severity of the previous winter as indicated by the maximum ice cover in the Baltic. The density of larvae was independent of larval density of the previous year suggesting that gobiid production primarily depends on the spawning stock size and may hence be regulated by density-independent factors.     

Population and size-specific distribution of Atlantic salmon Salmo salar in the Baltic Sea over five decades

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population. However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea.     

Effect of ammonium load on the growth of attached microalgae in the northern Baltic Sea

The effect of ammonium discharge from a food factory on the growth of attached microalgae was monitored north of the Hanko peninsula, on the southwestern coast of Finland. The impact of the discharge was studied at twelve localities, at four stages of seasonal succession. The microalgae were sampled from glass slides exposed at 0.4 m depth for two weeks. The variables measured for the microalgal growth were chlorophylla, primary production and total organic carbon (TOC). These were compared with planktonic chlorophylla and nutrient concentrations. The growth of attached microalgae displayed a consistent pattern of spatial distribution. Depending on season, TOC and primary production values were 7 to 70 times higher and chlorophylla values up to 1000 times higher close to the effluent outlet than in undisturbed areas of the archipelago. The microalgal samples near the discharge were characterized by low TOC/chlorophylla and TOC/primary production ratios. The temporal consistency of microalgal distribution illustrates the advantages of using attached algal assemblages in monitoring programmes.     

Impacts of nutrient enrichment and sediment on phytoplankton community structure in the northern Baltic Sea

A three-week mesocosm experiment was conducted in order to study the effects of bottom sediment and nutrient enrichment on phytoplankton and zooplankton community structure in the Archipelago Sea, northern Baltic Sea. The transparent polyethylene enclosures included the whole water column and varied in volume from 30 to 40 m³. There were two types of enclosures: some with natural sediment as a bottom and others with a plastic bottom. The experiment was a 2 × 2 factorial design with presence of sediment and nutrient enrichment as treatment factors. Both the sediment presence and nutrient enrichment significantly increased water nutrient concentrations and the rate of primary production. However, external nutrient enrichment and the presence of sediment stimulated the growth of different phytoplankton groups, indicating that the effect of sediment was not related to nutrient fluxes alone, but involved more complex interactions. External nutrient enrichment was primarily channelled to picoplanktonic cyanobacteria, the biomass of which increased four- to fivefold due to enrichment. The presence of sediment increased the biomass of cryptophytes, chrysophytes and prasinophytes, but decreased the biomass of N₂-fixing cyanobacteria. Zooplankton biomass increased during the experiment, but was not affected by the treatments. The study shows that sediment plays a significant role in phytoplankton dynamics, underlining the importance of including sediment in shallow-water mesocosm experiments. Handling editor: J. Padisak     

The chemical character and burial of phosphorus in shallow coastal sediments in the northeastern Baltic Sea

The chemical composition and vertical distribution of sediment phosphorus (P) in shallow coastal sediments of the northeastern Baltic Sea (BS) were characterized by sequential extraction. Different P forms were related to chemical and physical properties of the sediments and the chemistry of pore water and near-bottom water. Sediment P composition varied among the sampling sites located in the Archipelago Sea (AS) and along the northern coast of the Gulf of Finland (GoF): the organic rich sites were high in organic P (OP), while apatite-P dominated in the area affected by sediment transportation. Although the near-bottom water was oxic, the sediments released P. Release of P was most pronounced at the site with high sediment OP and reduced conditions in the sediment-water interface, indicating that P had its origins in organic sources as well as in reducible iron (Fe) oxyhydroxides. The results suggest that even though these coastal areas are shallow enough to lack salinity stratification typical for the brackish BS, they are vulnerable to seasonal oxygen (O₂) depletion and P release because of their patchy bottom topography, which restricts mixing of water. Furthermore, coastal basins accumulate organic matter (OM) and OP, degradation of which further diminishes O₂ and creates the potential for P release from the sediment. In these conditions, an abundance of labile OP may cause marked efflux of P from sediment reserves in the long-term.     

Long‐term temporal and spatial trends in eutrophication status of the Baltic Sea

Much of the Baltic Sea is currently classified as ‘affected by eutrophication’. The causes for this are twofold. First, current levels of nutrient inputs (nitrogen and phosphorus) from human activities exceed the natural processing capacity with an accumulation of nutrients in the Baltic Sea over the last 50–100 years. Secondly, the Baltic Sea is naturally susceptible to nutrient enrichment due to a combination of long retention times and stratification restricting ventilation of deep waters. Here, based on a unique data set collated from research activities and long‐term monitoring programs, we report on the temporal and spatial trends of eutrophication status for the open Baltic Sea over a 112‐year period using the HELCOM Eutrophication Assessment Tool (HEAT 3.0). Further, we analyse variation in the confidence of the eutrophication status assessment based on a systematic quantitative approach using coefficients of variation in the observations. The classifications in our assessment indicate that the first signs of eutrophication emerged in the mid‐1950s and the central parts of the Baltic Sea changed from being unaffected by eutrophication to being affected. We document improvements in eutrophication status that are direct consequences of long‐term efforts to reduce the inputs of nutrients. The reductions in both nitrogen and phosphorus loads have led to large‐scale alleviation of eutrophication and to a healthier Baltic Sea. Reduced confidence in our assessment is seen more recently due to reductions in the scope of monitoring programs. Our study sets a baseline for implementation of the ecosystem‐based management strategies and policies currently in place including the EU Marine Strategy Framework Directives and the HELCOM Baltic Sea Action Plan.     

Seasonal variability in the individual carbon content of the calanoid copepod Acartia bifilosa from the northern Baltic Sea

The seasonal variation in the carbon content of the calanoid copepod Acartia bifilosa was studied in the northern Baltic Sea. Monthly length-weight relationships were followed from November 1990 to October 1991 by analysing the organic carbon content of individuals, using a high temperature combustion method. The monthly length (L) on carbon (C) regressions of copepodites were best described by power functions (C = aL^b), where 75% to 88% of the variation could be explained by length. Nauplii length explained less of their carbon content (63–71 %). The carbon content per length was highest in summer when the nutritional situation was good and temperature at its highest. However, the carbon-length regressions differed significantly between months, except in winter, when two subsequent months had similar regressions. The results pointed out the importance of seasonal variation as well as the risk of making errors if biomass estimates based on length on carbon regressions are used. Carbon analysis should, if possible, be done on every sample or should at least be tested as to whether the regressions are consistent with the study material before estimations are calculated.     

Specific gravity and vertical distribution of sprat eggs in the Baltic Sea

During peak spawning of sprat Sprattus sprattus in the Baltic Sea in May–June egg specific gravity averaged ± s . d . 1·00858 ± 0·00116 g cm⁻³ but was significantly higher in the beginning and significantly lower towards the end of the spawning season. A close relationship was found between egg diameter and egg specific gravity ( r ² = 0·71). This relationship, however, changed during the spawning season indicating that some other factor was involved causing the decrease in specific gravity during the spawning period. The vertical egg distribution changed during the spawning season: eggs were distributed mainly in the deep layers early in the season, occurred in and above the permanent halocline during peak spawning, and above the halocline towards the end of the spawning season. Consequently, poor oxygen conditions in the deep layers and low temperatures in layers between the halocline and the developing thermocline may affect egg development. Thus, opportunities for egg development vary over the spawning season and among spawning areas, and depending on frequency of saline water inflows into the Baltic Sea and severity of winters, between years.     

Phytoplankton community responses to nutrient and iron enrichment under different nitrogen to phosphorus ratios in the northern Baltic Sea

The impact of nutrient enrichment on the phytoplankton community structure, and particularly cyanobacteria, was studied in a 3-week mesocosm experiment conducted in August 2001 in the Archipelago Sea, a part of the northern Baltic Sea. The factorial design experiment included daily additions of nitrogen (N) and phosphorus (P) at two mass ratios, 1N:1P and 7N:1P, respectively, additions of iron (Fe) and a synthetic chelator, ethylenediaminetetraacetic acid (EDTA). The floating enclosures (400 l) were sampled for analyses of phytoplankton biomass and community structure, phytoplankton primary production, chlorophyll a, nutrients, and hepatotoxins. Chlorophyll a concentration, phytoplankton biomass and primary production increased most in the 7N:1P treatment. The increase was mainly due to an abundant growth of chlorophytes (Dictyosphaerium subsolitarium, Kirchneriella spp., Monoraphidium contortum, and Oocystis spp.), pennate diatoms (especially Nitzschia spp.), dinophytes and the chroococcalean cyanobacterium Synechococcus sp. The nutrient enrichments had no effect on the total biomass of N₂-fixing cyanobacteria. Nevertheless, the biomass of Anabaena spp. was highest in the enrichments with a low N/P ratio. Chlorophyll a concentration and total phytoplankton biomass were not affected by Fe or EDTA, but Fe alone had a positive effect on the chlorophyte Kirchneriella sp. The N₂-fixing cyanobacteria Aphanizomenon sp. responded positively to Fe alone and to both Fe and EDTA added together. The hepatotoxin concentration increased during the experiment, but no clear responses to nutrient enrichments were found. Our study showed species-specific responses to nutrient enrichments among the N₂-fixing cyanobacteria. Although the total phytoplankton production was not Fe-limited; the availability of Fe clearly affected the phytoplankton community structure.     

Variation in fat content and fatty‐acid composition of the Baltic herring Clupea harengus membras

The fat content and fatty‐acid profiles of herring, Clupea harengus membras, from the southern Baltic Sea varied depending on when (fishing season) and where (fishing grounds) the fish were caught as well as on their size and sex. The fat, protein and dry matter content and the fatty‐acid profiles were assayed in C. h. membras muscle tissue. The changes observed in fatty‐acid profiles were determined by factors such as specimen mass and fat content, which, in turn, depended on fishing season. This is explained by dietary differences between juvenile and older fish. Gonad maturation and spawning in the latter are also factors. The study results provide confirmation of the hypothesis that polyunsaturated fatty acids (PUFA), in particular docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA), play vital roles in the sexual maturation of C. h. membras.     

Physiological response of the copepod Pseudocalanus sp. in the Baltic Sea at different thermal scenarios

We studied the physiological response of Pseudocalanus sp. under four different temperature elevation regimes: +0, +2, +4 and +6 °C above the decadal average temperature in the Western Baltic Sea. We measured fecal pellet (FP) production rates, which was taken as a proxy of ingestion, egg production (EPR) and respiration rates. Experiments lasted from mid‐February to end April, corresponding most of the observations to the postspring bloom phase. We combined small scale incubations with the use of big (ca. 1400 L) mesocosms, which have previously been shown to be appropriate when studying phyto‐ and zooplankton succession, and the water used for the incubations was taken from the mesocosm tanks. Given that the phytoplankton succession varied between the four thermal scenarios, we evaluated (excepting in the case of the respiration rates, where incubations were carried out using 0.2 μm filtered water) both the temperature and the associated food concentration effects. Respiration and ingestion rates were found to increase with temperature. As for EPR, they also increased with temperature during the bloom, but remained at low and constant values during the postbloom in all the four treatments due to the food limitation. Linked to the temperature rise, we also detected an increase in instantaneous mortality rates and a reduction in the net growth efficiency. Finally, we discuss the potential implications of our findings for the spring phyto‐ and zooplankton succession under the forecasted climate warming, as well as for the fisheries in the Baltic Sea, where Pseudocalanus sp. is a key species.     

Distributions,life cycles and hybridization of two Mysis relicta group species (Crustacea: Mysida) in the northern Baltic Sea and Lake Båven

We used electrophoretically identified material to assess the geographical distributions, life cycles and interspecific hybridization of two sibling species of the Mysis relicta species group (sp. I and sp. II) in the northern Baltic region. In the Gulf of Finland, sp. I prevails in inshore waters and sp. II in the open sea; the distributions overlap in the outer archipelago zone. In the Gulf of Bothnia, only sp. II was found in the southern part (Bothnian Sea), whereas the two species coexist throughout the northerly Bothnian Bay. Both the local and large-scale distributions are salinity-related, but salinity alone does not explain the differences. The two species exhibit different patterns of geographical variation in their life histories. In strict sympatry in the north they have identical two-year life cycles with winter breeding. Further south (Gulf of Finland), sp. I exhibits a predominantly one-year winter-breeding cycle, whereas sp. II breeds throughout the year. The patterns comply with the concept of a great phenotypic flexibility and environmental control of life history characteristics in the Mysis relicta group, and make a contrast to the stable life cycle of the congeneric M. mixta. F1 hybrids between the two M. relicta group species were found at a low frequency (0.6%) in the Bothnian Bay, but not in other areas of sympatry. This revised version was published online in July 2006 with corrections to the Cover Date.