Report of A Survey of the State of the Environment in the Estonia, Latvia and Lithuania

In connection with the change in economic principles and disintegration of big industrial enterprises, the environmental pollution in Baltic States decreased considerably in 1991-1996. Although there are still many things waiting to be done, the improvement of the Baltic environmental situation during the years of independence gives us a cause to be optimistic.     

Risk Analysis of Hunting of Seal Populations in the Baltic

Vulnerabilities of grey seal ( Halichoerus grypus) and ringed seal ( Phoca hispida) populations in the Baltic Sea were evaluated for potential opening of the populations for hunting. We used ecological risk analysis to assess the effects of environmental and demographic stochasticity and uncertain and partly missing population data on the modeled outcomes. The impact of different harvesting strategies on the long-term sustainability of seal populations was analyzed with four different models with increasing complexity and population detail. It appears the simpler the population model used, the more overconfident results it gave with regard to the hunting policy to be adopted. Therefore, it proves risky in population management decisions to rely on simplistic calculations based on growth rate and estimated population size alone. This is even more so if the population estimates have a wide error margin. Due to an unknown, but presumably positive number of seal kills in the Baltic at present, the sustainable harvest is likely to be close to zero for both seal species. Our risk analysis strongly suggests refraining from Baltic seal hunting now, with their current population sizes, and in the future if the development of population numbers cannot be assessed accurately enough.     

Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil,the priapulid Halicryptus spinulosus

Genetic variation of 16 allozyme loci in 397 Halicryptus spinulosus (Priapulida) revealed overall polymorphism of P=0.438 and Hardy-Weinberg expectations for heterozygosity of H _e=0.060 for Baltic Sea stocks, H _e=0.143 for the White Sea and H _e=0.121 for Iceland. Maximal unbiased standard distances of D=0.0693 separated Baltic and White Sea populations. Nordic and Baltic populations could be distinguished by allozymes, but Baltic subsamples proved cohesive. Gene flow amounted to effective exchange values per generation of N _m=2.94 over 650 km of continuous habitat, N _m=10.65 over 175 km, and N _m=13.85 over 20 km. Gene flow started to decrease with geographic distance beyond a dispersal threshold of 20 km, but hierarchical G _ST-statistics indicated light isolation by distance beyond a minimum of 8 km. Gene flow is high for a benthic worm assumed to lack dispersal by pelagic larva, a paradox which cannot be resolved now. Baltic populations are characterized by lower heterozygosity than Nordic stocks. In the Baltic Basin, temporally continuous brackish-water conditions have only existed for the past 7000 years. The two possible colonization routes of H. spinulosus to the geologically young Baltic Sea imply genetic drift, whether by founder effect (sweepstake colonization from Iceland) or by refugial bottlenecking during the Ancylus phase of the Baltic Basin after a direct connection to the White Sea had been sequestered. Continued genetic drift is emphasized by lower heterozygosity in the ecologically unstable Belt Sea compared to the central Baltic. Allozymes falsify the reduced-mutability hypothesis to explain bradytelic evolution of Priapulida. Regional genetic homogeneity, ample polymorphism, and preference for anoxic black mud qualify H. spinulosus populations as indicators of microevolutionary responses to water circulation regimes or pollution in the Baltic Sea.     

The role of environmental and spatial factors for the composition of aquatic bacterial communities

This study investigates the importance of local vs. spatial factors on bacterial community composition of 35 rock pools at the Baltic Sea coast. The pools were located in five distinct spatial clusters over a total scale of <500 m and differed widely in terms of water chemistry. To determine the fractions of the variance in bacterial community composition (BCC) between rock pools that are explained by local environmental vs. spatial factors, a variance partitioning procedure using partial canonical correspondence analysis was performed. Three environmental variables (salinity, chlorophyll a concentration, and water color) had a significant effect on BCC, irrespective of the spatial location of the pools. Vice versa, there was a significant effect of spatial factors on BCC irrespective of any of the environmental factors included in this study. Hence, the patchy spatial distribution of the pools was partly reflected in the composition of the bacterial communities in the pools, which might be caused by congruent colonization events of adjacent pools, such as simultaneous seaspray inputs or direct exchange of bacteria via connecting rivulets. This study shows that the composition of planktonic bacteria can show provincialism at small spatial scales, which is likely to be caused by environmental conditions as well as historical events.     

Are there two species of the polychaete genus Marenzelleria in Europe?

North Sea and Baltic Sea populations of the introduced polychaete Marenzelleria viridis (Verrill, 1873) reproduce at different times (spring and autumn, respectively). Enzyme separation by starch gel electrophoresis revealed major differences between specimens from the Baltic Sea and those from the North Sea (collected in 1992 and 1993) but a high degree of homogeneity among populations from the same sea. Three enzyme loci, glucose-6-phosphate isomerase (GPI-A, GPI-B) and malate dehydrogenase (MDH), were fixed to 100% by different alleles in the North and Baltic Sea populations, respectively. Different alleles are dominant for mitochondrial aspartate aminotransferase (mAAT) with allele frequencies of ca. 0.97 in all sampled populations from the North Sea and Baltic Sea, respectively, but heterozygotes were found in all populations. These genetic differences could be due to environmentally induced selection or genetically different origins of the populations, suggesting that populations of the genus Marenzelleria in the North and Baltic Seas may be two different species.     

Photosynthetic production and its regulating factors in the Baltic Sea

Measurements of primary production in the Baltic Sea over a period of 2 years (1969/1971) by the ¹⁴C method are reported. The rate of total annual production for the Baltic Sea proper and the Gulf of Finland including coastal areas appears to lie between 35 and 40 gC·m², a rate which can be assumed as characteristic for oligotrophic waters with a low production rate. An apparent increase in productivity between the 2 years could be noted. Calculations for the water mass below the halocline show that between 0.7 and 0.8 ml/l oxygen is used up during a period of 3 months required for the development of anoxic conditions. The major portion of the organic matter in the Baltic Sea undergoes decomposition in the sub-photic zone above the halocline. The ratio of organic material in deep water to the total available oxidizable matter appears to remain constant with time; therefore, the apparent increase in productivity in the photic layer does not affect apprecibly the development of anoxic conditions in deep water. Of the different areas investigated, the Gulf of Finland seems to be the most productive. In the southern Baltic Sea, the rate of production over 5 years between 1966 and 1971 has not changed appreciably. In fact, it shows a trend toward a slight decrease, whereas, in other areas of the Baltic Sea, the rate of production indicates an apparent increase. Comparison with oceanic coastal areas of the west coast of Sweden shows that the rate of production there is higher than in the Baltic Sea. Of the micronutrients controlling productivity in the photic layer of the Baltic Sea, nitrate has a stronger limiting effect on plant growth than phosphate, a fact which is in agreement with existing conceptions on the subject. Dissolved iron does not seem to exert any appreciable influence on the productivity.     

Self-purification capacity and management of Baltic coastal ecosystems

It is expected that the Baltic region becomes a major centre of economic growth and prosperity in Europe already during this decade (Anon. 2000). Therefore, an Agenda 21 for the Baltic region (Baltic 21) was developed to ensure a sustainable development. Especially the coastal ecosystems are subject to increasing anthropogenic pressure e.g. eutrophication, traffic, harbours, tourism or offshore wind parks. Eutrophication remains the main ecological problem in the Baltic Sea and has serious negative social and economical consequences. Inner and outer coastal waters play an important role as buffers and filters for the Baltic proper. Consequently, the utilization and preservation of their self-purification capacity is of great importance. Combined results of our own coastal research and of the international workshop ‘Baltic coastal ecosystems: structure, function and coastal zone management’ (Rostock University, November 2001) are presented here. Conclusions for an improved integrated coastal zone management of Baltic coastal ecosystems will be presented.     

Colonization of Europe by two American genetic types or species of the genus Marenzelleria (Polychaeta: Spionidae). An electrophoretic analysis of allozymes

Allozyme electrophoresis was conducted in an attempt to identify the origin ofMarenzelleria sp. found in the North Sea and Baltic Sea. The analysis covered eight enzymes with ten loti from nine populations found on the North American Atlantic toast, these populations in the North Sea and five populations in the Baltic. The North Sea spionids correspond to the Type IMarenzelleria from North American coastal waters between Barnstable Harbor (Massachusetts) and Cape Henlopen (Delaware). Nei's genetic distance between these North American populations and those from the North Sea wasD = 0.010 to 0.020. TheMarenzelleria sp. found in the Baltic Sea very probably stems from North American populations of Type II found from the region of Chesapeake Bay (Trippe Bay) south to the Currituck Sound (North Carolina). The genetic distance between these North American populations and the Baltic populations isD = 0.000 to 0.001. The invaders appear to have lost little of their genetic variation while colonizing the North and Baltic Seas. Probably, both colonizing events tan be attributed to large numbers of individuals reaching Europe simultaneously on one or more occasions. In addition, aMarenzelleria Type III was found by electrophoresis among specimens from Currituck Sound (North Carolina), rohere it is sympatric withMarenzelleria Type II. Salinity is discussed as an important factor for the establishment ofMarenzelleria Type I in the North Sea and Type II in the Baltic Sea.     

Chemical composition of Leptocythere psammophila (Crustacea: Ostracoda) as influenced by winter metabolism and summer supplies

Leptocythere psammophila is well known for the ecophenotypic variations of the ornamentation of its carapace. To test the respective influences of the environmental parameters on its ornamentation, 41 living specimens were collected in the Baltic Sea, the North Sea and in the English Channel during spring, summer and winter (April 1987, September and December 1988). The carbonate carapaces of these specimens were analyzed by means of electron microprobe. Thirteen elements were detected. Statistical comparisons of means and variances were performed using classical F and t-tests. Data were submitted to Normalized Principal Component Analysis and Discriminant Analysis. There are no significant differences between chemical composition of the carapaces from the different stations. However, the variability of Baltic Sea specimens is lower than that of other stations. Nonetheless, strong and significant differences appear when samples are gathered by season. The chemical composition of summer individuals is controlled by variations in water salinity and in fine grain terrigenous sediment supply. The composition of winter samples is related to the incorporation of Mg. In the case of spring specimens, both these factors have to be considered. Discriminant analysis between winter and non-winter samples sets correctly 84% of the individuals. The wrongly classified specimens are interpreted as ostracods that did not molt during the season of sampling. By attributing them to their possible season of molting, the result of the Normalized Principal Component Analysis improves. Differences in ornamentation are slight. They occur between the samples from the Baltic Sea and those from the other stations. The former have larger and less numerous punctations and lack the smooth surface in the anteroventral part of the carapace. The punctation diameter is more variable during summer than during spring. In the range of environments investigated, ornamentation of the carapace of L. psammophila seems unaffected by the seasonal environmental variations whereas its composition exhibits strong differences.     

Predator-inducible changes in blue mussels from the predator-free Baltic Sea

Blue mussels, Mytilus spp., have inhabited the brackish Baltic Sea, an environment lacking predatory crabs and starfish, for several thousand years. In this paper we examined whether Baltic Mytilus that were transplanted to the North Sea showed predator-inducible plasticity like their "marine" conspecifics. Our experiments showed that native North Sea Mytilus changed their morphology when exposed to waterborne scents from shore-crabs and starfish. These predators induced different kinds of changes, with emphasis on shell thickness and adductor muscle size, respectively. Baltic Mytilus responded in a similar way to crab scents, whereas starfish scents had a relatively weak effect on the morphology. Crab and starfish scent induced strengthening of the byssal attachment in North Sea Mytilus, with crabs providing more stimulation than starfish. Baltic mussels also improved the byssal attachment when exposed to either of the predators, but the attachment strength, as well as the response to crabs, were relatively weaker than that of North Sea mussels. We conclude that inducible plasticity still is present in Baltic Mytilus, despite their recent evolution in a predator-free environment. There is probably no strong selective pressure against inducible plasticity, but it could also be maintained in the population by gene flow from Mytilus in the adjoining North Sea. The question whether Baltic Mytilus are M. edulis or M. trossulus may also be relevant for the present results.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

Radioactivity in the Baltic Sea

The Baltic Sea is, like the Mediterranean, a marginal sea, which with the Black Sea, were marine environments contaminated from the Chernobyl accident.

Radiocaesium and plutonium isotopes were studied in water, sediment and macroalgae in the Baltic Sea since 1982. the inventory of ¹³⁷Cs in the Baltic increased from 0.65 PBq to 5.85 PBq following the Chernobyl accident. the corresponding increase for ^{239 + 240}Pu was less significant and yielded 1.5 TBq to a total value of 16.5 TBq.

For plutonium, 98% is trapped in the sediment and the net-exchange of this element through the Baltic straits is very small (1 GBq/year), while for radiocaesium, 45% is in the water phase and there is a net-loss of 60 000 GBq annually into the adjacent water (Kattegatt).     

Functional divergence in heat shock response following rapid speciation of Fucus spp. in the Baltic Sea

In the Baltic Sea, the broadly distributed brown alga Fucus vesiculosus coexists in sympatry over part of its range (south west Gulf of Bothnia) with the Baltic endemic F. radicans sp. nov, while further north in colder and lower-salinity areas of the Baltic F. radicans occurs alone (north west Gulf of Bothnia). F. radicans appears to have arisen via rapid speciation from F. vesiculosus within the recent history of the Baltic (ca. 7500 BP). Possible functional divergence between the two species was investigated by comparing stress-responsive gene expression in a common-garden experiment. The experiment used two allopatric populations of Fucus vesiculosus from the Skagerrak (North Sea) and Central Baltic, as well as F. radicans from the same Central Baltic site. The two species in sympatry displayed divergent heat shock responses, while F. vesiculosus populations from allopatric sites did not. F. radicans was more sensitive to heat shock at 25°C, either alone or together with high irradiance and desiccation, than Baltic or Skagerrak F. vesiculosus. The results indicate that rapid functional divergence in the inducible heat shock response has occurred between sympatric species on a timescale of thousands of years.     

Benthic diversity gradients and shifting baselines: implications for assessing environmental status

The increasing pressure on marine biodiversity emphasizes the importance of finding benchmarks against which to assess change. This is, however, a notoriously difficult task in estuarine ecosystems, where environmental gradients are steep, and where benthic biodiversity is highly variable in space and time. Although recent emphasis on diverse, healthy benthic communities in legislative frameworks has increased the number of indices developed for assessing benthic status, there is a lack of quantitative baselines in benthic diversity that would enable comparisons across broad spatial scales, encompassing different environmental settings and bioregions. By taking advantage of long-term monitoring data, spanning hundreds of stations over the past 40 years, we provide a comprehensive analysis of benthic a, beta, and gamma diversity, encompassing the entire' salinity gradient of the open sea areas of the large, brackish-water Baltic Sea. Using a relatively simple measure, average regional diversity, we define area-specific reference conditions and acceptable deviation against which to gauge current conditions in benthic macrofaunal diversity. Results show a severely impaired condition throughout large areas of the Baltic for the assessment period 2001-2006. All ecosystems are plagued by baselines that shift in time and space, and their definition is not trivial, but average regional diversity may offer a transparent way to deal with such changes in low-diversity systems. Identifying baselines will be of increasing importance given the potential of climatic drivers to interact with local anthropogenic stressors to affect patterns of biodiversity. Our analysis provides an evaluation of the current condition in a system that has been heavily influenced by anthropogenic impact and changing oceanographic conditions, and it provides a basis for future impact assessment and ecosystem-based management.     

Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

Large-scale diversity and biogeography of benthic copepods in European waters

A large-scale database concerning benthic copepods from the Arctic, Baltic Sea, North Sea, British Isles, Adriatic Sea and Crete was compiled to assess species richness, biodiversity, communities, ecological range size and biogeographical patterns. The Adriatic showed the highest evenness and the most species-rich communities. Assemblages from the North Sea, British Isles, Baltic and Crete had a lower evenness. The British Isles were characterised by impoverished communities. The ecological specificity of copepod species showed two diverging trends: higher specificity of species in more diverse assemblages was observed in the Adriatic, North Sea and Baltic. A uniformly high species specificity disregarding sample diversity was found on Crete and in the British Isles. Benthic copepod communities showed distinct patterns that clearly fit the predefined geographical regions. Communities were distinguishable and β-diversity was found to be high around Europe, indicating a high species turnover on the scale of this investigation. The British Isles and the North Sea were found to be faunistic links to the Baltic and the Arctic.     

Temporal and spatial variation in the growth rates of Baltic herring (<Emphasis Type="Italic">Clupea harengus membras</Emphasis> L.) larvae during summer

It has been suggested that larval survival determines the year-class strength in most marine fish species. During their growth and development, the ability of the larvae to catch prey and avoid predation will increase. However, the factors affecting short-term changes in the growth of Baltic Sea herring have been little studied in the field. We collected Baltic herring (Clupea harengus membras L.) larvae from five different towing areas in the Archipelago Sea (SW Finland) during May and June 1989, right after the main spawning season. Twenty thousand two hundred and ten larvae were analysed and the area-specific growth rate (i.e. increase in standard length) was estimated by tracing the larval cohorts from the length-frequency data. This represents the first Baltic herring study with daily sampling during a long study period. The growth rate was related to environmental factors, such as temperature, number of zooplankters, and wind speed and direction. Large variation in larval growth rate occurred between areas: lowest and highest growth rates were 0.18 and 0.52 mm·day^-1. Temperature was an important variable controlling larval-fish growth rate. An increase of one 1°C in average water temperature corresponded to an increase in growth rate of 0.043 mm·day^-1. This may have been caused either by a direct temperature effect (changes in metabolic rate) or by the indirect effect of changes in food availability. We also found the densest herring populations in the areas with highest average water temperature. However, temperature and larval growth rate both increased towards the inner archipelago.     

New indicator species in the Baltic zooplankton in 1972

The occurrence of zooplankton species, indicating a salinewater influx from the North Sea into the Baltic Sea in 1972, is described. Particular attention is paid to 8 species so far unknown in this region (Ectopleura dumortieri, Rosacea plicata, Tomopteris kefersteini, Aetideus armatus, Centropages typicus, Eucalanus elongatus, Metridia lucens, Electra pilosa). The appearance of these zooplankters in the Southern Baltic Sea is discussed in relation to the hhdrographic changes taking place in this area in 1972. Based on these data, assessments are made regarding the intensity of dynamic exchange processes in the Baltic and North Atlantic waters in 1972.     

Effects of salinity on spermatozoa motility,percentage of fertilized eggs and egg development of Baltic cod (Gadus morhua), and implications for cod stock fluctuations in the Baltic

Variation in cod stock strength in the Baltic Sea is considered to be dependent on abiotic conditions such as salinity and water oxygen content in the spawning areas (the Baltic deep basins). Spawning cod were caught off northern Gotland, Sweden, from April to June in 1989 and 1990. Our investigation revealed a reduction in spermatozoan motility and a subsequent reduction in the percentage of fertilized eggs in salinities of 10 to 12. Normal egg development required a minimum salinity of 11. This coincides with the osmolality of the seminal plasma and egg yolk; i.e., the Baltic cod is adapted to hyperosmotic conditions for spawning and is thus totally dependent on periodical inflows of saline water from the North Sea.     

Environmental cues and constraints affecting the seasonality of dominant calanoid copepods in brackish, coastal waters: a case study of Acartia, Temora and Eurytemora species in the south-west Baltic

Information on physiological rates and tolerances helps one gain a cause-and-effect understanding of the role that some environmental (bottom–up) factors play in regulating the seasonality and productivity of key species. We combined the results of laboratory experiments on reproductive success and field time series data on adult abundance to explore factors controlling the seasonality of Acartia spp., Eurytemora affinis and Temora longicornis, key copepods of brackish, coastal and temperate environments. Patterns in laboratory and field data were discussed using a metabolic framework that included the effects of ‘controlling’, ‘masking’ and ‘directive’ environmental factors. Over a 5-year period, changes in adult abundance within two south-west Baltic field sites (Kiel Fjord Pier, 54°19′89N, 10°09′06E, 12–21 psu, and North/Baltic Sea Canal NOK, 54°20′45N, 9°57′02E, 4–10 psu) were evaluated with respect to changes in temperature, salinity, day length and chlorophyll a concentration. Acartia spp. dominated the copepod assemblage at both sites (up to 16,764 and 21,771 females m^?3 at NOK and Pier) and was 4 to 10 times more abundant than E. affinis (to 2,939?m^?3 at NOK) and T. longicornis (to 1,959?m^?3 at Pier), respectively. Species-specific salinity tolerance explains differences in adult abundance between sampling sites whereas phenological differences among species are best explained by the influence of species-specific thermal windows and prey requirements supporting survival and egg production. Multiple intrinsic and extrinsic (environmental) factors influence the production of different egg types (normal and resting), regulate life-history strategies and influence match–mismatch dynamics.