Dynamics of phytoplankton in the brackish-water inlet Pojoviken,southern coast of Finland

The water exchange between the brackish-water firth Pojoviken and the Baltic Sea is restricted by a shallow sill (6 m). An outflowing, oligohaline surface layer is isolated from the nutrient-rich mesohaline deep water by a pycnocline at a depth of 6–10 m. During the ice-free period phytoplankton production is chiefly regulated by the river discharge regime. Contrary to the situation in the outer archipelago and the sea zone, in Pojoviken phytoplankton production continues until late autumn, because the stable salinity stratification prevents the phytoplankton from sinking below the critical depth for production. The phytoplankton composition seems to be regulated chiefly by salinity. The salinity interval 2–2.5 is apparently the critical range where brackish-water phytoplankton changes to an assemblage composed of typical freshwater species.     

Where did Marenzelleria spp. (Polychaeta: Spionidae) in Europe come from?

Abundant populations of Marenzelleria spp. were reported for the first time in the North Sea during the late 1970s and then in the Baltic Sea in 1985. Genetic analysis by means of allozyme electrophoresis and sequencing of a segment of mitochondrial 16srDNA showed that two different genetic types or sibling species of Marenzelleria were present in Europe. Marenzelleria Type I is found only in the North Sea, whereas Type II has been found in both the North Sea and the Baltic Sea. The North Sea animals (Type I) correspond to Type I specimens found in North American coastal waters between Nova Scotia and Cape Henlopen (Delaware), while Marenzelleria Type II from the Baltic Sea correspond to Marenzelleria type II animals from the Arctic (Tuktoyaktuk Harbor, Northwest Territories, Canada), New Hampshire and coastal waters between Chesapeake Bay southward to the Ogeechee River (Georgia). Human mediated introduction (by shipping) and natural range expansion are discussed as the possible causes of these virtually simultaneous invasions by two sibling species. Marenzelleria Type I appears to colonize habitats with a higher salinity and/or in which salinities tend to fluctuate considerably. The osmolality of the coelomic fluid after acclimation to various salinities between 0.25 and 18 is the same for both sibling species/genetic types. Although the two types do not differ in respect of hyperosmoregulation (<10), differences may exist in their cell volume regulation or its time course in the almost isoosmotic range at salinities >10.     

Diversity and abundance of “Pelagibacterales” (SAR11) in the Baltic Sea salinity gradient

The candidate order “Pelagibacterales” (SAR11) is one of the most abundant bacterial orders in ocean surface waters and, periodically, in freshwater lakes. The presence of several stable phylogenetic lineages comprising “Pelagibacterales” correlates with the physico-chemical parameters in aquatic environments. A previous amplicon sequencing study covering the bacterial community in the salinity gradient of the Baltic Sea suggested that pelagibacteral subclade SAR11-I was replaced by SAR11-IIIa in the mesohaline region of the Baltic Sea. In this current study, we investigated the cellular abundances of “Pelagibacterales” subclades along the Baltic Sea salinity gradient using catalyzed reporter deposition fluorescence in situ hybridization (CARD-FISH). The results obtained with a newly designed probe, which exclusively detected SAR11-IIIa, were compared to CARD-FISH abundances of the marine SAR11-I/II subclade and the freshwater lineage SAR11-IIIb (LD12). The results showed that SAR11-IIIa was abundant in oligohaline–mesohaline conditions (salinities 2.7–13.3), with maximal abundances at a salinity of 7 (up to 35% of total Bacteria, quantified with a universal bacterial probe EUB). As expected, SAR11-I/II was abundant (27% of EUB) in the marine parts of the Baltic Sea, whereas counts of the freshwater lineage SAR11-IIIb were below the detection limit at all stations. The shift from SAR11-IIIa to SAR11-I/II was confirmed in the vertical salinity gradient in the deeper basins of the Baltic Sea. These findings were consistent with an overlapping but defined distribution of SAR11-I/II and SAR11-IIIa in the salinity gradient of the Baltic Sea and suggested the adaptation of SAR11-IIIa for growth and survival in mesohaline conditions.     

Molecular identification key based on PCR/RFLP for three polychaete sibling species of the genus <Emphasis Type="Italic">Marenzelleria</Emphasis>, and the species’ current distribution in the Baltic Sea

Studies of Marenzelleria species were often hampered by identification uncertainties when using morphological characters only. A newly developed PCR/RFLP protocol allows a more efficient discrimination of the three species Marenzelleria viridis, Marenzelleria neglecta and Marenzelleria arctia currently known for the Baltic Sea. The protocol is based on PCR amplification of two mitochondrial DNA gene segments (16S, COI) followed by digestion with restriction enzymes. As it is faster and cheaper than PCR/sequencing protocols used so far, the protocol is recommended for large-scale analyses. The markers allow an undoubted determination of species irrespective of life stage or condition of the worms in the samples. The protocol was validated on about 950 specimens sampled at more than 30 sites of the Baltic and the North Sea, and on specimens from populations of the North American east coast. Besides this test we used mitochondrial DNA sequences (16S, COI, Cytb) and starch gel electrophoresis to further investigate the distribution of the three Marenzelleria species in the Baltic Sea. The results show that M. viridis (formerly genetic type I or M. cf. wireni) occurred in the Öresund area, in the south western as well as in the eastern Baltic Sea, where it is found sympatric with M. neglecta. Allozyme electrophoresis indicated an introduction by range expansion from the North Sea. The second species, M. arctia, was only found in the northern Baltic Sea, where it sometimes occurred sympatric with M. neglecta or M. viridis. For Baltic M. arctia, the most probable way of introduction is by ship ballast water from the European Arctic. There is an urgent need for a new genetic analysis of all Marenzelleria populations of the Baltic Sea to unravel the current distribution of the three species.     

Marenzelleria cf. viridis (Polychaeta: Spionidae) – ecophysiological adaptations to a life in the coastal waters of the Baltic Sea

Since its introduction into the Baltic Sea about ten years ago, the polychaete species Marenzelleria cf. viridis has spread rapidly. Meanwhile, this spionid settled in remarkable numbers predominantly in the coastal waters which provide some of the more variable and unstable habitats in terms of abiotic conditions. In the present paper, some of the underlying biochemical and physiological processes were experimentally examined which enables this polychaete to deal with several kinds of environmental stress such as variations in salinities, low oxygen concentrations and occurrence of hydrogen sulphide. The results obtained reveal that in the process of acclimation to different salinities free amino acids are involved, mainly alanine and glycine. The content of these amino acids were increased in worms acclimated to a higher salinity. When exposed to low ambient oxygen concentrations (severe hypoxia) M. cf. viridis started to produce energy anaerobically via pathways known from several other marine organisms. The presence of hydrogen sulphide in higher concentrations (1 mmol l^-1) results in a more rapid production of succinate (indicator for anaerobic energy production via the succinate-propionate pathway) as compared to hypoxia alone, indicating a higher energy demand. Nevertheless, when exposed to enhanced ambient hydrogen sulphide concentrations (3 mmol l^-1), M. cf. viridis is less affected by this toxic compound than other marine species, such as the related species Marenzelleria cf. wireni (North Sea). The field data indicate that the metabolic response obtained under experimental conditions is also important in the worm's natural habitat. Although the extent of anaerobic metabolites produced as well as the amount of thiosulphate (main sulphide detoxification product) was lower in the field as compared to the experiments, M. cf. viridis very likely has to face hypoxia and hydrogen sulphide in higher concentrations in its natural habitat.     

Salinity gradient of the Baltic Sea limits the reproduction and population expansion of the newly invaded comb jelly Mnemiopsis leidyi

The recent invasion of the comb jelly Mnemiopsis leidyi into northern European waters is of major public and scientific concern. One of the key features making M. leidyi a successful invader is its high fecundity combined with fast growth rates. However, little is known about physiological limitations to its reproduction and consequent possible abiotic restrictions to its dispersal. To evaluate the invasion potential of M. leidyi into the brackish Baltic Sea we studied in situ egg production rates in different regions and at different salinities in the laboratory, representing the salinity gradient of the Baltic Sea. During October 2009 M. leidyi actively reproduced over large areas of the Baltic Sea. Egg production rates scaled with animal size but decreased significantly with decreasing salinity, both in the field (7-29) and in laboratory experiments (6-33). Temperature and zooplankton, i.e. food abundance, could not explain the observed differences. Reproduction rates at conditions representing the Kattegat, south western and central Baltic Sea, respectively, were 2.8 fold higher at the highest salinities (33 and 25) than at intermediate salinities (10 and 15) and 21 times higher compared from intermediate to the lowest salinity tested (6). Higher salinity areas such as the Kattegat, and to a lower extent the south western Baltic, seem to act as source regions for the M. leidyi population in the central Baltic Sea where a self-sustaining population, due to the low salinity, cannot be maintained.     

Comparative morphology of larvae of coastal crabs (Crustacea: Decapoda: Varunidae)

The larval development of three crabs of the Varunidae family, (Hemigrapsus sanguineus, H. penicillatus, and H. longitarsis), widely spread in Russian waters of the Sea of Japan, were studied under laboratory conditions. At a temperature of 20–22°C and a salinity of 32‰ about 30% of larvae a complete developmental cycle, including five zoeal stages and megalopa, took from 22 to 30 days. All larval stages are illustrated and described in detail. Zoea I and zoea II of the studied crabs are not distinguishable. Zoea III–V of these species differ in the number of dorsomedial setae on the abdominal somite I and in the number of setae on the posterodorsal arch. The megalopae of three Hemigrapsus species possess a different number of segments, aesthetascs and setae on the antennular exopod. In spite of the great similarity of larvae of genera Hemigrapsus and Eriocheir, the latter possesses a number of distinctive features in all developmental stages, supporting the separation of these genera.     

Water salinity effects on embryogenesis of the lesser sandeel,Ammodytes tobianus (Linnaeus, 1758)

Effects of water salinity (3.0–9.0 PSU) on the duration and outcome of embryogenesis in the lesser sandeel (Ammodytes tobianus (L.) as well as on survival, size, and malformation rate of the newly hatched larvae were followed, based on observations involving equipment consisting of a microscope and still and digital cameras. The images obtained and measurements of egg and larval dimensions were analyzed using the NIS elements Br. computer software package. The lowest salinity tested (3.0 PSU), similar to that in the inshore Baltic Sea waters native for the species studied, produced a lower fertilization rate and resulted in extended embryogenesis, reduced embryo survival, changes in the hatch size, and increased larval malformation rate. The high-end salinity (7.5–9.0 PSU), higher than that prevailing in the species’s spawning grounds in the Baltic Sea, resulted — similarly to the low-end salinity — in disturbed development and growth of the embryos.     

Feeding biology of the pelagic larvae of Marenzelleria cf. viridis (Polychaeta: Spionidae) from the Baltic Sea

Phytoplankton < 20 µm was a principal dietary component of the larvae of Marenzelleria cf. viridis. Maximum ingested particle size increased as animal size increased, reaching a maximum diameter of 80 µm for larvae with 6 to 10 setigers. The larvae started ingesting particulate matter at the 1-setiger stage and were able selectively to ingest phytoplankton and polystyrene particles of various sizes. Larvae in the 6 to 10-setiger size group did not differ from those in the 11 to 17-setiger size group in respect of size selectivity for polystyrene particles. The gut passage time for Chlorella vulgaris was 20 min. The ingestion rate was limited by food concentrations even at concentrations much higher than those encountered in the natural biotope, saturation being reached at a concentration of 28.5 times 10⁶ cells ml^-1 (117.7 mg C l^-1. The low maximum filtration rate of only 1.19 µl ind.^-1 h^-1 indicates that the filtering capacity of the larvae is low. The larvae are still capable of food uptake at 1 °C. Further experiments demonstrated that larval growth and survival were strongly dependent on both food concentration and quality. Larval growth was food-limited under biotope conditions of the Darss–Zingst Boddens and even more so under Baltic Sea conditions. The results indicate that Marenzelleria cf. viridis is a species adapted to eutrophic conditions prevailing in brackish waters.     

Survival, activity and feeding ability of Baltic cod (Gadus morhua) yolk-sac larvae at different salinities

Due to unfavourable conditions (declines in salinity and water oxygen content) in the spawning areas, there has been a considerable decrease in the Baltic cod stock since the beginning of the 1980s, and consequently a decrease in catches. In order to examine the feasibility of introducing yolk-sac larvae in areas of low salinity to improve the stock, laboratory experiments were performed on the effects of salinity on the survival, level of activity and feeding ability of larvae. Yolk-sac larvae from spawning cod caught off northern Gotland, Sweden, were exposed to four different salinities: 10 and 15%○ (salinities of the main spawning areas); and 5 and 7%○ (salinities in the Bothnian Sea and the Baltic proper respectively).
The survival of yolk-sac larvae was high at all salinities, even though there was an indication of higher mortality at low salinities in less viable larval groups. No differences were found in swimming speed or feeding ability at the four salinities, but a significant difference in vertical distribution was recorded. There were significant differences in survival, vertical distribution and feeding ability among larval groups, which indicates that larval quality or viability is of greater importance for larval survival than salinity, in the range of 5–15%○.     

Kombinierte Wirkungen von Temperatur,Salzgehalt und Cadmium auf Entwicklung,Wachstum und Mortalität der Larven vonMytilus edulis aus der westlichen Ostsee

The response surface method was used to study the effects of temperature, salinity and Cd contamination on the development of fertilized eggs ofMytilus edulis from the western Baltic Sea to the veliger stage as well as on growth and cumulative mortality of a veliger population. The reactions observed differ considerably with reference to temperature and salinity. This is discussed in respect to the mode of life of the larvae and to the environmental conditions of the western Baltic Sea. Cd only slightly influences various temperature-dependent life functions, but strongly modifies those depending on salinity. The development optimum is shifted to higher salinities with increasing Cd concentrations of the medium, while that of survival and growth is shifted to lower salinities. These factor interactions modify the tolerance limits. In addition, the rates of factor interactions on the larval stages change with the degree of development. The trochophora stage proves to be most sensitive to the factors studied. A significant influence of cadmium on different life functions is found from concentrations of about 50 ppb on.     

Brackish-water submergence of the common periwinkle,Littorina littorea,and its digenean parasites in the Baltic Sea and in the Kattegat

North Sea and Baltic Sea populations ofLittorina littorea differ with respect to their vertical distribution. In the North SeaL. littorea is strictly intertidal while in the Baltic Sea maximum population densities occur in the sublittoral. Levels of infestation with larval digenetic trematodes diminish qualitatively (number of species recorded) and quantitatively (number of hosts infested) with decreasing salinity. Both the host and two parasite species —Cryptocotyle lingua andMicrophallus pygmaeus — display brackish-water submergence under conditions of reduced surface salinity.     

A broad transition zone between an inner Baltic hybrid swarm and a pure North Sea subspecies of Macoma balthica (Mollusca, Bivalvia)

The populations of the bivalve clam Macoma balthica in the low-salinity Northern Baltic Sea represent an admixture of two strongly diverged genomic origins, the Pacific Macoma balthica balthica (approx. 60% genomic contribution) and Atlantic Macoma balthica rubra (40%). Using allozyme and mtDNA characters, we describe the broad transition from this hybrid swarm to the pure M. b. rubra in the saline North Sea waters, spanning hundreds of kilometre distance. The zone is centred in the strong salinity gradient of the narrow Öresund strait and in the adjacent Western Baltic. Yet the multilocus clines show no simple and smoothly monotonic gradation: they involve local reversals and strong differences between neighbouring populations. The transitions in different characters are not strictly coincident, and the extent of introgression varies among loci. The Atlantic influence extends further into the Baltic in samples from the southern and eastern Baltic coasts than on the western coast, and further in deeper bottoms than at shallow (< 1 m) sites. This fits with the counterclockwise net circulation pattern and with a presumably weaker salinity barrier for invading Atlantic type larvae in saline deeper water, and corresponding facilitation of outwards drift of Baltic larvae in diluted surface waters. Genotypic disequilibria were strong particularly in the shallow-water samples of the steepest transition zone. This suggests larval mixing from different sources and limited interbreeding in that area, which makes a stark contrast to the evidence of thorough amalgamation of the distinct genomic origins in the inner Baltic hybrid swarm of equilibrium structure.     

Ascarophis sp. (Nematoda: Spirurida) attaining sexual maturity inGammarus spp. (Crustacea)

Ascarophis sp., including sexually mature adult worms, was commonly recorded in the amphipodGammarus oceanicus Segerstrle in the northern Baltic Sea and also inGammarus sp. in estuarine localities in the New Brunswick region of the north-western Atlantic. Species of the genusAscarophis van Beneden (Nematoda: Cystidicolidae) are known as parasites of marine and brackish water fishes, whereas generally only larval forms have been reported from crustaceans. Adults as well as larval stages are described (LM and SEM) and the infection dynamics is analysed in relation to the amphipod population. The results suggest a direct transmission of embryonated eggs to new amphipods, although this remains to be verified experimentally.     

Reproductive traits of the cold-seep symbiotic mussel Idas modiolaeformis: gametogenesis and larval biology

We describe the first reproductive features of a chemosynthetic mussel collected at cold seeps from the eastern Mediterranean Sea. Idas modiolaeformis (Bivalvia, Mytilidae) is a hermaphroditic species in which production of male and female gametes likely alternates, a feature regarded as an adaptation to patchy and ephemeral habitats. By using fluorescent in situ hybridization, we demonstrate that bacterial symbionts, while present within the gills, are absent within acini that enclose female gametes and male gametes. This supports the hypothesis of environmental acquisition of symbionts in chemosynthetic mytilids. Prodissoconch I (PI) is relatively small compared to prodissoconch II (PII), suggesting a planktotrophic larval stage. Diameters of the two larval shells are in the range of sizes reported for mytilids, with a PII size between that of the shallow Mytilus edulis and that of the cold-seep mussel "Bathymodiolus" childressi.     

Prorocentrum minimum (Dinophyceae) in the Baltic Sea: morphology, occurrence—a review

The potentially toxic dinoflagellate Prorocentrum minimum (Pavillard) Schiller has successfully established in the Baltic Sea in the last two decades. A review of the invasion history is presented as well as new data on the spatial and inter-annual variability of this species and its relation to salinity, temperature, and nutrient concentrations. A short literature review of the morphological characters of the Baltic P. minimum is also included.From 1993 to 2002, P. minimum was a regular component of the summer and autumn plankton flora of the Baltic Sea proper and the Gulf of Finland. Its abundance varied considerably inter-annually and did not show any clear trends during the period. Abundance of P. minimum was significantly higher in the nutrient-enriched Bay of Mecklenburg (German coast) and the southern Baltic proper than in the central and northern Baltic proper and the Gulf of Finland, where its abundance was mostly sparse. In coastal waters P. minimum occasionally reached densities of several million cells per litre and dominated phytoplankton biomass (>90%).Abundance of the Baltic P. minimum was generally not related to salinity or temperature. It could be a dominant species at both high and low salinity (over 15 and 4.8 PSU), and its temperature range was broad (from 2.7 to 26.4 °C). However, dense populations usually occurred from July to October at temperatures above 10 °C.Further, there appears to be a positive correlation between the success of P. minimum in the Baltic Sea and high concentrations of total phosphorus and nitrogen.This tolerant and morphologically variable dinoflagellate seems to be a morphospecies without subtaxa, which can expand its range in the Baltic Sea, especially in nutrient-rich coastal waters.     

An environmental gradient dominates ecological and genetic differentiation of marine invertebrates between the North and Baltic Sea

Environmental gradients have emerged as important barriers to structuring populations and species distributions. We set out to test whether the strong salinity gradient from the marine North Sea to the brackish Baltic Sea in northern Europe represents an ecological and genetic break, and to identify life history traits that correlate with the strength of this break. We accumulated mitochondrial cytochrome oxidase subunit 1 sequence data, and data on the distribution, salinity tolerance, and life history for 28 species belonging to the Cnidaria, Crustacea, Echinodermata, Mollusca, Polychaeta, and Gastrotricha. We included seven non‐native species covering a broad range of times since introduction, in order to gain insight into the pace of adaptation and differentiation. We calculated measures of genetic diversity and differentiation across the environmental gradient, coalescent times, and migration rates between North and Baltic Sea populations, and analyzed correlations between genetic and life history data. The majority of investigated species is either genetically differentiated and/or adapted to the lower salinity conditions of the Baltic Sea. Species exhibiting population structure have a range of patterns of genetic diversity in comparison with the North Sea, from lower in the Baltic Sea to higher in the Baltic Sea, or equally diverse in North and Baltic Sea. Two of the non‐native species showed signs of genetic differentiation, their times since introduction to the Baltic Sea being about 80 and >700 years, respectively. Our results indicate that the transition from North Sea to Baltic Sea represents a genetic and ecological break: The diversity of genetic patterns points toward independent trajectories in the Baltic compared with the North Sea, and ecological differences with regard to salinity tolerance are common. The North Sea–Baltic Sea region provides a unique setting to study evolutionary adaptation during colonization processes at different stages by jointly considering native and non‐native species.     

Chinese mitten crab <Emphasis Type="Italic">Eriocheir sinensis</Emphasis> in the Baltic Sea—a supply-side invader?

Although the Chinese mitten crab Eriocheir sinensis (H. Milne-Edwards, 1853) (Crustacea, Decapoda, Varunidae) invaded the Baltic Sea about 80 years ago, published information on its present distribution and abundance in this region is lacking. We provide here information on its Baltic-wide distribution and long-term population dynamics. The species has been found all over the coastal Baltic Sea and also in some adjacent rivers and lakes. The Chinese mitten crab appears to have increased in abundance in recent years in the northeastern part of the Baltic Sea (Gulf of Finland, Gulf of Riga, northern Baltic Proper). Higher catch rates were observed in spring (April–June) and autumn (September–November). The size variation of crabs in different samples was low (mean carapace width 6.1–6.3 cm). Despite findings of gravid females, the reproduction of the mitten crab in the central, northern and eastern Baltic region is considered unlikely due to low salinity and the individuals caught are assumed to actively migrate into the region from the species’ main European distribution area (southeastern North Sea), certainly over 1500 km migration distance. Thus, the dynamics of the North Sea population is probably regulating, at least in part, the occurrence of the Chinese mitten crab in the Baltic Sea area.     

The lifecycle of <Emphasis Type="Italic">Dichelyne minutus</Emphasis> (Rudolphi, 1819) (Nematoda: Cucullanidae) in the estuarine biocenosis of the Black Sea

The lifecycle, the host–parasite system, and the ecological features of the nematode Dichelyne minutus (Rudolphi, 1819), which parasitizes invertebrates and fish in the estuarine biocenosis located at the influx of the Chornaya River into the Black Sea (off Sevastopol), have been studied. The host–parasite system of D. minutus includes the polychaete Hediste diversicolor Müller, 1776 (as an obligatory intermediate host) and nine fish species, of which seven are definitive hosts and two are accidental or captive hosts. It has been found that the lifecycle of D. minutus in the biocoenosis of the Black Sea differs from the lifecycle of this nematode that inhabits the Baltic and North seas. In the studied biocoenosis, nematode larvae occur in polychaetes and fish only in the spring and summer; no larvae are found in the autumn (the study was not conducted in the winter). The nematode parasitizes the polychaete H. diversicolor in the spring; the main source of infection in this period is obviously nematode eggs that were laid in the autumn and have overwintered in the environment. The infection process ends by early summer. The seasonal and size–age dynamics of nematode infection of the round goby, Neogobius melanostomus (Pallas, 1814), are analyzed taking the specifics of fish biology into account. The short period of infection, as characterized by the active emission of nematode larvae, their low survival in polychaetes and fish, a short lifecycle and the mortality of mature nematodes after egg-laying in the autumn result in an over-scattered distribution (mostly of the negative-binomial type) of D. minutus in populations of all the hosts.