GENETIC AND MORPHOLOGICAL IDENTIFICATION OF FUCUS RADICANS SP. NOV. (FUCALES,PHAEOPHYCEAE) IN THE BRACKISH BALTIC SEA1

Brown seaweeds of the genus Fucus occupy a wide variety of temperate coastal habitats. The genus is evolutionary dynamic with recent radiations to form morphologically distinct taxa. In the brackish Baltic Sea, fucoids are the only perennial canopy‐forming macroalgae. The most northern populations of Fucus occur permanently submerged in extremely low salinity (3–5 psu). These are currently referred to as Fucus vesiculosus L. but are morphologically distinct with a narrow frond without bladders. We report here that a population of this unique morphotype is reproductively isolated from a truly sympatric population of common F. vesiculosus and conclude that the northern morphotype represents a previously undescribed species. We describe Fucus radicans sp. nov., which is attached and dioecious with broadly elliptic receptacles, characterized by a richly branched narrow flat frond (2–5 mm), short thallus (<26 cm), and a high capacity for vegetative recruitment of attached plants. Analysis of five highly polymorphic microsatellite DNA loci showed genetic differentiation between sympatric populations of F. radicans and F. vesiculosus, whereas allopatric populations of the same species revealed a coherent pattern of genetic variation. Sequences of the RUBISCO region in F. radicans were identical to or differing at only one to two dinucleotide positions from those of F. vesiculosus, indicating a recent common origin of the two species.     

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.     

Thiamin dynamics during the adult life cycle of Atlantic salmon (Salmo salar)

Thiamin is an essential water-soluble B vitamin known for its wide range of metabolic functions and antioxidant properties. Over the past decades, reproductive failures induced by thiamin deficiency have been observed in several salmonid species worldwide, but it is unclear why this micronutrient deficiency arises. Few studies have compared thiamin concentrations in systems of salmonid populations with or without documented thiamin deficiency. Moreover, it is not well known whether and how thiamin concentration changes during the marine feeding phase and the spawning migration. Therefore, samples of Atlantic salmon (Salmo salar) were collected when actively feeding in the open Baltic Sea, after the sea migration to natal rivers, after river migration, and during the spawning period. To compare populations of Baltic salmon with systems without documented thiamin deficiency, a population of landlocked salmon located in Lake Vänern (Sweden) was sampled as well as salmon from Norwegian rivers draining into the North Atlantic Ocean. Results showed the highest mean thiamin concentrations in Lake Vänern salmon, followed by North Atlantic, and the lowest in Baltic populations. Therefore, salmon in the Baltic Sea seem to be consistently more constrained by thiamin than those in other systems. Condition factor and body length had little to no effect on thiamin concentrations in all systems, suggesting that there is no relation between the body condition of salmon and thiamin deficiency. In our large spatiotemporal comparison of salmon populations, thiamin concentrations declined toward spawning in all studied systems, suggesting that the reduction in thiamin concentration arises as a natural consequence of starvation rather than to be related to thiamin deficiency in the system. These results suggest that factors affecting accumulation during the marine feeding phase are key for understanding the thiamin deficiency in salmonids.     

Small-scale spatial and temporal variation in the demographic processes underlying the large-scale decline of eiders in the Baltic Sea

The application of uniform conservation schemes often fails to account for small-scale spatial variation in the drivers of population decline. Demographic comparisons of imperilled populations across locations are therefore crucial for successful conservation, but progress is hampered by lack of long-term data from more than a single population. The recent large-scale decline of eider ducks (Somateria mollissima) in the Baltic Sea is ideal for determining to what extent mechanisms underlying population decline can be extrapolated over larger areas. We utilized stochastic demographic methods incorporating both environmental and sampling variation to assess small-scale spatial and temporal variation in the population dynamics of eiders at Söderskär (eastern range-margin) and Tvärminne (core breeding area), situated 130 km apart. The stochastic growth rate models accurately predicted the observed differences in the rate of decline between sites and time periods. At Söderskär, established breeder survival had by far the greatest elasticity, whereas elasticity was more evenly distributed among vital rates at Tvärminne. Although the study sites showed the single largest difference in fecundity, stochastic life table response experiment analyses revealed that reduced adult female survival at Tvärminne mainly determined the observed difference in growth rates between sites. In contrast, reduced fecundity primarily differentiated the past population increase from the present population decline at Söderskär. Our results demonstrate that different mechanisms may be associated with population decline across adjacent geographic locations, and indicate that dispersal of first-time breeders may be important for population dynamics. Safeguarding adult female survival and/or fecundity should be prioritized in management efforts.     

Habitat traits and food availability determine the response of marine invertebrates to ocean acidification

Energy availability and local adaptation are major components in mediating the effects of ocean acidification (OA) on marine species. In a long‐term study, we investigated the effects of food availability and elevated pCO₂ (ca. 400, 1000 and 3000 μatm) on growth of newly settled Amphibalanus (Balanus) improvisus to reproduction, and on their offspring. We also compared two different populations, which were presumed to differ in their sensitivity to pCO₂ due to differing habitat conditions: Kiel Fjord, Germany (Western Baltic Sea) with naturally strong pCO₂ fluctuations, and the Tjärnö Archipelago, Sweden (Skagerrak) with far lower fluctuations. Over 20 weeks, survival, growth, reproduction and shell strength of Kiel barnacles were all unaffected by elevated pCO₂, regardless of food availability. Moulting frequency and shell corrosion increased with increasing pCO₂ in adults. Larval development and juvenile growth of the F1 generation were tolerant to increased pCO₂, irrespective of parental treatment. In contrast, elevated pCO₂ had a strong negative impact on survival of Tjärnö barnacles. Specimens from this population were able to withstand moderate levels of elevated pCO₂ over 5 weeks when food was plentiful but showed reduced growth under food limitation. Severe levels of elevated pCO₂ negatively impacted growth of Tjärnö barnacles in both food treatments. We demonstrate a conspicuously higher tolerance to elevated pCO₂ in Kiel barnacles than in Tjärnö barnacles. This tolerance was carried over from adults to their offspring. Our findings indicate that populations from fluctuating pCO₂ environments are more tolerant to elevated pCO₂ than populations from more stable pCO₂ habitats. We furthermore provide evidence that energy availability can mediate the ability of barnacles to withstand moderate CO₂ stress. Considering the high tolerance of Kiel specimens and the possibility to adapt over many generations, near future OA alone does not seem to present a major threat for A. improvisus.     

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.     

Morphological and genetic variation of Fucus in the eastern Gulf of Bothnia,northern Baltic Sea

Fucus vesiculosus and F. radicans (Phaeophyceae) are important habitat-formers on rocky shores in the Bothnian Sea. While both species occur sympatrically along the entire western Bothnian Sea coast, F. radicans has been found only in the northern part of the eastern coast. According to previous studies, the two species can be distinguished based on morphology, F. radicans having narrower thalli and a bushier appearance. However, marine mapping in the eastern Bothnian Sea has revealed that high morphological variation in Fucus, partly caused by gradients in salinity and exposure, makes differentiation between the two species difficult. We studied morphological and genetic variation to find out whether the two Fucus species can be differentiated in the south-eastern Bothnian Sea, and if F. radicans occurs in the area. The study was carried out in six subareas including 350 km of coast, with a salinity gradient of 3.5–6.5 PSU, and varying wave exposure. We found a gradual change towards smaller and narrower thalli and a higher number of holdfasts in Fucus populations when moving northwards to lower salinities. Distinct Fucus morphs were often found within the study sites but the morphs were genetically differentiated only at one study site in the Skaftung subarea, suggesting the occurrence of both species. However, in the Vasa subarea the sample size for analysing genetic differentiation was low due to high clonality. In the Luvia subarea south of Skaftung, Fucus morphology corresponded to that of F. radicans in earlier studies but the population was genetically more similar to F. vesiculosus in the southern subareas. We conclude that by using only morphological characteristics it is not possible to differentiate between the two species in central and northern parts of the eastern Bothnian Sea. Based on genetic analyses, the southernmost known occurrence of F. radicans in the eastern Bothnian Sea is in Skaftung.     

Unravelling the complexity of salt marsh ‘Fucus cottonii’ forms (Phaeophyceae,Fucales)

Genetic affiliation, nuclear DNA content, and gamete functioning were examined in small salt marsh Fucus from three localities in western Ireland. Individuals with small and dioecious receptacles were found at all localities, but production of germlings was only evident at Locality 1. Here, the Fucus vegetation formed a morphological cline from F. vesiculosus with bladders in the mid-intertidal to small Fucus individuals lacking bladders in the salt marsh of the upper intertidal. Measurements of nuclear DNA content ranged from 1–1.8 pg at this locality, with F. vesiculosus individuals in the lower range. At the two other localities, the small salt marsh Fucus consisted of distinct morphological entities. Microsatellite analyses revealed that individuals at Locality 2 were derived mainly from F. vesiculosus, whereas those from Locality 3 were hybrids between F. vesiculosus and F. spiralis with greatest affiliation to F. spiralis. While the small salt marsh Fucus forms from Locality 2 had high nuclear DNA content (c. 4 pg) and were probably octoploids, the small salt marsh Fucus from Locality 3 formed two groups: one with high (3.9–4.6 pg) and one with low (1.5–1.9 pg) nuclear DNA content. Nuclear DNA content measured in individuals from Locality 3 varied between 1.1–2.8 pg in F. vesiculosus and 2–3.5 pg in F. spiralis, and showed a more or less stepwise increase in both species, consistent with polyploidy. We hypothesize that the small salt marsh Fucus forms originate from genome size changes in the parental taxa.     

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.     

Evidence of a hybrid-zone in Atlantic cod (Gadus morhua) in the Baltic and the Danish Belt Sea revealed by individual admixture analysis

The study of hybrid zones is central to our understanding of the genetic basis of reproductive isolation and speciation, yet very little is known about the extent and significance of hybrid zones in marine fishes. We examined the population structure of cod in the transition area between the North Sea and the Baltic Sea employing nine microsatellite loci. Genetic differentiation between the North Sea sample and the rest increased along a transect to the Baltic proper, with a large increase in level of differentiation occurring in the Western Baltic area. Our objective was to determine whether this pattern was caused purely by varying degrees of mechanical mixing of North Sea and Baltic Sea cod or by interbreeding and formation of a hybrid swarm. Simulation studies revealed that traditional Hardy-Weinberg analysis did not have sufficient power for detection of a Wahlund effect. However, using a model-based clustering method for individual admixture analysis, we were able to demonstrate the existence of intermediate genotypes in all samples from the transition area. Accordingly, our data were explained best by a model of a hybrid swarm flanked by pure nonadmixed populations in the North Sea and the Baltic Sea proper. Significant correlation of gene identities across loci (gametic phase disequilibrium) was found only in a sample from the Western Baltic, suggesting this area as the centre of the apparent hybrid zone. A hybrid zone for cod in the ecotone between the high-saline North Sea and the low-saline Baltic Sea is discussed in relation to its possible origin and maintenance, and in relation to a classical study of haemoglobin variation in cod from the Baltic Sea/Danish Belt Sea, suggesting mixing of two divergent populations without interbreeding.     

Differences in Salinity Tolerance and Gene Expression Between Two Populations of Atlantic Cod (Gadus morhua) in Response to Salinity Stress

Populations of marine fish, even from contrasting habitats, generally show low genetic differentiation at neutral genetic markers. Nevertheless, there is increasing evidence for differences in gene expression among populations that may be ascribed to adaptive divergence. Studying variation in salinity tolerance and gene expression among Atlantic cod (Gadus morhua) from two populations distributed across a steep salinity gradient, we observed high mortality (45% North Sea cod and 80% Baltic Sea cod) in a reciprocal common garden setup. Quantitative RT-PCR assays for expression of hsp70 and Na/K-ATPase α genes demonstrated significant differences in gene regulation within and between populations and treatment groups despite low sample sizes. Most interesting are the significant differences observed in expression of the Na/K-ATPase α gene in gill tissue between North Sea and Baltic cod. The findings strongly suggest that Atlantic cod are adapted to local saline conditions, despite relatively low levels of neutral genetic divergence between populations.     

Genetic population structure of turbot (Scophthalmus maximus L.) supports the presence of multiple hybrid zones for marine fishes in the transition zone between the Baltic Sea and the North Sea

Genetic population structure of turbot (Scophthalmus maximus L.) in the Northeast Atlantic was investigated using eight highly variable microsatellite loci. In total 706 individuals from eight locations with temporal replicates were assayed, covering an area from the French Bay of Biscay to the Aaland archipelago in the Baltic Sea. In contrast to previous genetic studies of turbot, we found significant genetic differentiation among samples with a maximum pairwise FST of 0.032. Limited or no genetic differentiation was found among samples within the Atlantic/North Sea area and within the Baltic Sea, suggesting high gene flow among populations in these areas. In contrast, there was a sharp cline in genetic differentiation going from the low saline Baltic Sea to the high saline North Sea. The data were explained best by two divergent populations connected by a hybrid zone; however, a mechanical mixing model could not be ruled out. A significant part of the genetic variance could be ascribed to variation among years within locality. Nevertheless, the population structure was relatively stable over time, suggesting that the observed pattern of genetic differentiation is biologically significant. This study suggests that hybrid zones are a common phenomenon for marine fishes in the transition area between the North Sea and the Baltic Sea and highlights the importance of using interspecific comparisons for inferring population structure in high gene flow species such as most marine fishes.     

Diversity of luciferase sequences and bioluminescence production in Baltic Sea Alexandrium ostenfeldii

The toxic dinoflagellate Alexandrium ostenfeldii is the only bioluminescent bloom-forming phytoplankton in coastal waters of the Baltic Sea. We analysed partial luciferase gene (lcf) sequences and bioluminescence production in Baltic A. ostenfeldii bloom populations to assess the distribution and consistency of the trait in the Baltic Sea, and to evaluate applications for early detection of toxic blooms. Lcf was consistently present in 61 Baltic Sea A. ostenfeldii strains isolated from six separate bloom sites. All Baltic Sea strains except one produced bioluminescence. In contrast, the presence of lcf and the ability to produce bioluminescence did vary among strains from other parts of Europe. In phylogenetic analyses, lcf sequences of Baltic Sea strains clustered separately from North Sea strains, but variation between Baltic Sea strains was not sufficient to distinguish between bloom populations. Clustering of the lcf marker was similar to internal transcribed spacer (ITS) sequences with differences being minor and limited to the lowest hierarchical clusters, indicating a similar rate of evolution of the two genes. In relation to monitoring, the consistent presence of lcf and close coupling of lcf with bioluminescence suggests that bioluminescence can be used to reliably monitor toxic bloom-forming A. ostenfeldii in the Baltic Sea.     

GENETIC STRUCTURE IN POPULATIONS OF FUCUS VESICULOSUS (PHAEOPHYCEAE) OVER SPATIAL SCALES FROM 10 M TO 800 KM1

Recent studies showing consequences of species’ genetic diversity on ecosystem performance raise the concern of how key ecosystem species are genetically structured. The bladder wrack Fucus vesiculosus L. is a dominant species of macroalga in the northern Atlantic, and it is particularly important as a habitat‐forming species in the Baltic Sea. We examined the genetic structure of populations of F. vesiculosus with a hierarchical approach from a within‐shore scale (10 m) to a between‐seas scale (Baltic Sea–Skagerrak, 800 km). Analysis of five microsatellite loci showed that population differentiation was generally strong (average F_ST = 12%), being significant at all spatial scales investigated (10¹, 10³, 10^4–5, 10⁶ m). Genetic differentiation between seas (Baltic Sea and Skagerrak) was substantial. Nevertheless, the effects of isolation by distance were stronger within seas than between seas. Notably, Baltic summer‐reproducing populations showed a strong within‐sea, between‐area (70 km) genetic structure, while Baltic autumn‐reproducing populations and Skagerrak summer‐reproducing populations revealed most genetic diversity between samples within areas (<1 km). Despite such differences in overall structure, Baltic populations of summer‐ and autumn‐reproducing morphs did not separate in a cluster analysis, indicating minor, if any, barriers to gene flow between them. Our results have important implications for management and conservation of F. vesiculosus, and we raise a number of concerns about how genetic variability should be preserved within this species.     

Salinity gradient shapes distance decay of similarity among parasite communities in three marine fishes

Published data were used to compare the distance decay of similarity in parasite communities of three marine fish hosts: Atlantic cod Gadus morhua, the dab Limanda limanda and the flounder Platichthys flesus in two adjacent areas that differ with respect to the strength of a salinity gradient. In the Baltic Sea, which exhibits a strong salinity gradient from its connection with the North Sea in the west to its head in the north‐east, parasite communities in all three fish hosts showed a significant decline of similarity with increasing distance. In contrast, among host populations in the North Sea, which is a fully marine environment, there was no such decline or only a weak relationship. The results suggest that environmental gradients like salinity can be strong driving forces behind patterns of distance decay in parasite communities of fishes.     

FREQUENT CLONALITY IN FUCOIDS (FUCUS RADICANS AND FUCUS VESICULOSUS; FUCALES,PHAEOPHYCEAE) IN THE BALTIC SEA1

Asexual reproduction by cloning may affect the genetic structure of populations, their potential to evolve, and, among foundation species, contributions to ecosystem functions. Macroalgae of the genus Fucus are known to produce attached plants only by sexual recruitment. Recently, however, clones of attached plants recruited by asexual reproduction were observed in a few populations of Fucus radicans Bergström et L. Kautsky and F. vesiculosus L. inside the Baltic Sea. Herein we assess the distribution and prevalence of clonality in Baltic fucoids using nine polymorphic microsatellite loci and samples of F. radicans and F. vesiculosus from 13 Baltic sites. Clonality was more common in F. radicans than in F. vesiculosus, and in both species it tended to be most common in northern Baltic sites, although variation among close populations was sometimes extensive. Individual clonal lineages were mostly restricted to single or nearby locations, but one clonal lineage of F. radicans dominated five of 10 populations and was widely distributed over 550 × 100 km of coast. Populations dominated by a few clonal lineages were common in F. radicans, and these were less genetically variable than in other populations. As thalli recruited by cloning produced gametes, a possible explanation for this reduced genetic variation is that dominance of one or a few clonal lineages biases the gamete pool resulting in a decreased effective population size and thereby loss of genetic variation by genetic drift. Baltic fucoids are important habitat‐forming species, and genetic structure and presence of clonality have implications for conservation strategies.     

Phylogeographic evidence for the postglacial colonization of the North and Baltic Sea coasts from inland glacial refugia by Triglochin maritima L.

We investigated the geographical distribution of genetic variation in 67 individuals of Triglochin maritima from 38 localities across Europe using AFLP markers. Analysis of genetic variation resulted in the recognition of two major genetic groups. Apart from few geographical outliers, these are distributed (1) along the Atlantic coasts of Portugal, Spain and France and (2) in the North Sea area, the Baltic Sea area, at central European inland localities, the northern Adriatic Sea coast and the Mediterranean coast of southwest France. Considering possible range shifts of T. maritima in reaction to Quaternary climatic changes as deduced from the present-day northern temperature limit of the species, Quaternary changes of coastline in the North Sea area and the very recent origin of the Baltic Sea, we conclude that the coastal populations of T. maritima in the North Sea and Baltic Sea areas originated from inland populations.     

Local adaptation and oceanographic connectivity patterns explain genetic differentiation of a marine diatom across the North Sea–Baltic Sea salinity gradient

Drivers of population genetic structure are still poorly understood in marine micro‐organisms. We exploited the North Sea–Baltic Sea transition for investigating the seascape genetics of a marine diatom, Skeletonema marinoi. Eight polymorphic microsatellite loci were analysed in 354 individuals from ten locations to analyse population structure of the species along a 1500‐km‐long salinity gradient ranging from 3 to 30 psu. To test for salinity adaptation, salinity reaction norms were determined for sets of strains originating from three different salinity regimes of the gradient. Modelled oceanographic connectivity was compared to directional relative migration by correlation analyses to examine oceanographic drivers. Population genetic analyses showed distinct genetic divergence of a low‐salinity Baltic Sea population and a high‐salinity North Sea population, coinciding with the most evident physical dispersal barrier in the area, the Danish Straits. Baltic Sea populations displayed reduced genetic diversity compared to North Sea populations. Growth optima of low salinity isolates were significantly lower than those of strains from higher native salinities, indicating local salinity adaptation. Although the North Sea–Baltic Sea transition was identified as a barrier to gene flow, migration between Baltic Sea and North Sea populations occurred. However, the presence of differentiated neutral markers on each side of the transition zone suggests that migrants are maladapted. It is concluded that local salinity adaptation, supported by oceanographic connectivity patterns creating an asymmetric migration pattern between the Baltic Sea and the North Sea, determines genetic differentiation patterns in the transition zone.