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.     

Diversity of Aphanizomenon flos-aquae (cyanobacterium) populations along a Baltic Sea salinity gradient

Colony-forming cyanobacteria of the genus Aphanizomenon form massive blooms in the brackish water of the Baltic Sea during the warmest summer months. There have been recent suggestions claiming that the Baltic Sea Aphanizomenon species may be different from Aphanizomenon flos-aquae found in lakes. In this study, we examined variability in the morphology and 16S-23S rRNA internal transcribed spacer (ITS) sequences of A. flos-aquae populations along a salinity gradient from a string of lakes to a fjord-like extension of the Baltic Sea to the open Baltic Sea. Morphological differences among the populations were negligible. We found that the Baltic Sea was dominated (25 out of 27 sequences) by one ITS1-S (shorter band of ITS 1 [ITS1]) genotype, which also was found in the lakes. The lake populations of A. flos-aquae tended to be genetically more diverse than the Baltic Sea populations. Since the lake ITS1-S genotypes of A. flos-aquae are continuously introduced to the Baltic Sea via inflowing waters, it seems that only one ITS1 genotype is able to persist in the Baltic Sea populations. The results suggest that one of the ITS1-S genotypes found in the lakes is better adapted to the conditions of the Baltic Sea and that natural selection removes most of the lake genotypes from the Baltic Sea A. flos-aquae populations.     

Uptake of dissolved organic nitrogen by size-fractionated plankton along a salinity gradient from the North Sea to the Baltic Sea

The Baltic Sea is known for its ecological problems due to eutrophication caused by high nutrient input via nitrogen fixation and rivers, which deliver up to 70% of nitrogen in the form of dissolved organic nitrogen (DON) compounds. We therefore measured organic nitrogen uptake rates using self produced ¹⁵N labeled allochthonous (derived from Brassica napus and Phragmites sp.) and autochthonous (derived from Skeletonema costatum) DON at twelve stations along a salinity gradient (34 to 2) from the North Sea to the Baltic Sea in August/September 2009. Both labeled DON sources were exploited by the size fractions 0.2–1.6 μm (bacteria size fraction) and >1.6 μm (phytoplankton size fraction). Higher DON uptake rates were measured in the Baltic Sea compared to the North Sea, with rates of up to 1213 nmol N l^?1 h^?1. The autochthonous DON was the dominant nitrogen form used by the phytoplankton size fraction, whereas the heterotrophic bacteria size fraction preferred the allochthonous DON. We detected a moderate shift from >1.6 μm plankton dominated DON uptake in the North Sea and central Baltic Sea towards a 0.2–1.6 μm dominated DON uptake in the Bothnian Bay and a weak positive relationship between DON concentrations and uptake. These findings indicate that DON is an important component of plankton nutrition and can fuel primary production. It may therefore also contribute substantially to eutrophication in the Baltic Sea especially when inorganic nitrogen sources are depleted.     

Priority effects in a planktonic bloom-forming marine diatom

Priority effects occur when a species or genotype with earlier arrival has an advantage such that its relative abundance in the community or population is increased compared with later-arriving species. Few studies have dealt with this concept in the context of within-species competition. Skeletonema marinoi is a marine diatom that shows a high degree of genetic differentiation between populations over small geographical distances. To test whether historical events such as priority effects may have been important in inducing these patterns of population differentiation, we performed microcosm experiments with successive inoculation of different S. marinoi strains. Our results show that even in the absence of a numerical advantage, significant priority effects were evident. We propose that priority effects may be an important mechanism in initiating population genetic differentiation.     

Short-term photosynthetic responses in the diatom Nitzschia americana to a simulated salinity environment

The effect of short-term variations in saiinity on photosynthesisand respiration in an estuarine clone of the diatom Nitzschiaamericana was examined using a laboratory environmental simulationsystem. A computer-controlled culturing system simulated surfacelongitudinal salinity gradients in the Cape Fear River Estuary,NC by regulating the growth conditions of a continuous culturein real-time. Salinity changes were programmed based on a one-dimensionalhorizontal advection model of the Cape Fear River Estuary. Thesystem proved useful in evaluating phytoplanklon photosyntheticresponses to changes in its local environment on time scalessimilar to those operating in natural systems. As more sophisticatedmodels of the physical environment and particle transport areincorporated into the simulation program, realistic physiologicalmodels of phytoplankton production can be developed. Averagerates of net carbon fixation measured during the environmentalsimulation were compared to rates predicted by long-term adaptationstudies. These results indicate that rates of net carbon fixationdecline sharply following a rapid (hours) increase in salinity.Rates of net carbon fixation return, however, approximatelyto maximum predicted rates if the salinity environment is constantfor approximately 24 h, suggesting an adaptation period. Presumablythis time is necessary for N. americana to initiate physiologicalmechanisms responsible for osmoregulation and various compensatoryresponses to changes in salinity. These results further indicatethat significant errors may result when production models basedon long-term adaptive responses are used to describe phytoplanktonproductivity in variable environments.     

Genetic diversity of Saccharina latissima (Phaeophyceae) along a salinity gradient in the North Sea–Baltic Sea transition zone

The North Sea–Baltic Sea transition zone constitutes a boundary area for the kelp species Saccharina latissima due to a strong salinity gradient operating in the area. Furthermore, the existence of S. latissima there, along Danish waters, is fairly patchy as hard bottom is scarce. In this study, patterns of genetic diversity of S. latissima populations were evaluated along the salinity gradient area of Danish waters (here designated brackish) and were compared to reference sites (here designated marine) outside the gradient area, using microsatellite markers. The results showed that the S. latissima populations were structured into two clusters corresponding to brackish versus marine sites, and that gene flow was reduced both between clusters and between populations within clusters. In addition, results provided empirical evidence that marginal populations of S. latissima in the salinity gradient area exhibited a distinct genetic structure when compared to marine ones. Brackish populations were less diverse, more related, and showed increased differentiation over distance compared to marine populations. The isolation of the brackish S. latissima populations within the salinity gradient area of Danish waters in conjunction with their general low genetic diversity makes these populations vulnerable to ongoing environmental and climate change, predicted to result in declining salinity in the Baltic Sea area that may alter the future distribution and performance of S. latissima in the area.     

Biomass,composition and activity of organism assemblages along a salinity gradient in sea ice subjected to river discharge in the Baltic Sea

A study was undertaken to examine the activity and composition of the seasonal Baltic Sea land-fast sea-ice biota along a salinity gradient in March 2003 in a coastal location in the SW coast of Finland. Using a multi-variable data set, the less well-known algal and protozoan communities, and algal and bacterial production in relation to the physical and chemical environment were investigated. Also, the first coincident measurements of bacterial production and dissolved organic matter (DOM) in a sea-ice system are reported. Communities in sea ice were clearly autotrophy-dominated with algal biomass representing 79% of the total biomass. Protozoa and rotifers made up 18% of biomass in the ice and bacteria only 3%. Highest biomasses were found in mid-transect bottom ice. Water column assemblages were clearly more heterotrophic: 39% algae, 12% bacteria and 49% for rotifers and protozoa. Few significant correlations existed between DOM and bacterial variables, reflecting the complex origin of ice DOM. Dynamics of dissolved organic carbon, nitrogen and phosphorus (DOC, DON and DOP) were also uncoupled. A functional microbial loop is likely to be present in the studied ice. Existence of an under-ice freshwater plume affects the ecosystem functioning: Under-ice water communities are influenced directly by river-water mixing, whereas the ice system seems to be more independent—the interaction mainly taking place through the formation of active bottom communities.     

Rapid speciation in a newly opened postglacial marine environment, the Baltic Sea

Background  

Theory predicts that speciation can be quite rapid. Previous examples comprise a wide range of organisms such as sockeye salmon, polyploid hybrid plants, fruit flies and cichlid fishes. However, few studies have shown natural examples of rapid evolution giving rise to new species in marine environments.     

Geographic variation in fitness‐related traits of the bladderwrack Fucus vesiculosus along the Baltic Sea‐North Sea salinity gradient

In the course of the ongoing global intensification and diversification of human pressures, the study of variation patterns of biological traits along environmental gradients can provide relevant information on the performance of species under shifting conditions. The pronounced salinity gradient, co‐occurrence of multiple stressors, and accelerated rates of change make the Baltic Sea and its transition to North Sea a suitable region for this type of study. Focusing on the bladderwrack Fucus vesiculosus, one of the main foundation species on hard‐bottoms of the Baltic Sea, we analyzed the phenotypic variation among populations occurring along 2,000 km of coasts subjected to salinities from 4 to >30 and a variety of other stressors. Morphological and biochemical traits, including palatability for grazers, were recorded at 20 stations along the Baltic Sea and four stations in the North Sea. We evaluated in a common modeling framework the relative contribution of multiple environmental drivers to the observed trait patterns. Salinity was the main and, in some cases, the only environmental driver of the geographic trait variation in F. vesiculosus. The decrease in salinity from North Sea to Baltic Sea stations was accompanied by a decline in thallus size, photosynthetic pigments, and energy storage compounds, and affected the interaction of the alga with herbivores and epibiota. For some traits, drivers that vary locally such as wave exposure, light availability or nutrient enrichment were also important. The strong genetic population structure in this macroalgae might play a role in the generation and maintenance of phenotypic patterns across geographic scales. In light of our results, the desalination process projected for the Baltic Sea could have detrimental impacts on F. vesiculosus in areas close to its tolerance limit, affecting ecosystem functions such as habitat formation, primary production, and food supply.     

Electron requirements for carbon incorporation along a diel light cycle in three marine diatom species

Diatoms account for about 40% of primary production in highly productive ecosystems. The development of a new generation of fluorometers has made it possible to improve estimation of the electron transport rate from photosystem II, which, when coupled with the carbon incorporation rate enables estimation of the electrons required for carbon fixation. The aim of this study was to investigate the daily dynamics of these electron requirements as a function of the diel light cycle in three relevant diatom species and to apprehend if the method of estimating the electron transport rate can lead to different pictures of the dynamics. The results confirmed the species-dependent capacity for photoacclimation under increasing light levels. Despite daily variations in the photosynthetic parameters, the results of this study underline the low daily variability of the electron requirements estimated using functional absorption of the photosystem II compared to an estimation based on a specific absorption cross section of chlorophyll a. The stability of the electron requirements throughout the day would suggest it is potentially possible to estimate high-frequency primary production by using autonomous variable fluorescence measurements from ships-of-opportunity or moorings, without taking potential daily variation in this parameter into consideration, but this result has to be confirmed on natural phytoplankton assemblages. The results obtained in this study confirm the low electron requirements of diatoms to perform photosynthesis, and suggest a potential additional source of energy for carbon fixation, as recently described in the literature for this class.     

Diversity of Aphanizomenonflos-aquae (Cyanobacterium) Populations along a Baltic Sea Salinity Gradient

Colony-forming cyanobacteria of the genus Aphanizomenon form massive blooms in the brackish water of the Baltic Sea during the warmest summer months. There have been recent suggestions claiming that the Baltic Sea Aphanizomenon species may be different from Aphanizomenon flos-aquae found in lakes. In this study, we examined variability in the morphology and 16S-23S rRNA internal transcribed spacer (ITS) sequences of A. flos-aquae populations along a salinity gradient from a string of lakes to a fjord-like extension of the Baltic Sea to the open Baltic Sea. Morphological differences among the populations were negligible. We found that the Baltic Sea was dominated (25 out of 27 sequences) by one ITS1-S (shorter band of ITS 1 [ITS1]) genotype, which also was found in the lakes. The lake populations of A. flos-aquae tended to be genetically more diverse than the Baltic Sea populations. Since the lake ITS1-S genotypes of A. flos-aquae are continuously introduced to the Baltic Sea via inflowing waters, it seems that only one ITS1 genotype is able to persist in the Baltic Sea populations. The results suggest that one of the ITS1-S genotypes found in the lakes is better adapted to the conditions of the Baltic Sea and that natural selection removes most of the lake genotypes from the Baltic Sea A. flos-aquae populations.     

Anatomy of a cline: dissecting anti‐predatory adaptations in a marine gastropod along the U.S. Atlantic coast

The scope of anti‐predatory adaptation is expected to be greater in warm than in cold environments. High temperatures lower the costs associated with the production and maintenance of energetically expensive traits and enable ecological interactions to intensify. We tested this hypothesis by characterizing the expression of anti‐predatory morphology within a marine gastropod species (the knobbed whelk Busycon carica) over a large (> 1400 km) geographic area that spans more than 10°C annual temperature variation. We also conducted experimental predation studies with a powerful durophagous predator, the stone crab Menippe, to verify the anti‐predatory advantages of a heavily ornamented shell morphology (e.g. increased thickness, pronounced spines), and we used repair scar data to assess clinal variation in selective pressure from predators. We predicted that repair scar rates would be greatest in warm southernmost latitudes, and that expression of energetically costly anti‐predatory morphology would peak in concert with elevated predation pressures. Experiments confirmed that whelks with energetically costly, heavily ornamented shells had higher survivorship rates than those with weakly ornamented shells. As predicted, we also found that the expression of anti‐predatory traits was greatest in the southern part of B. carica's range. After standardizing shells for size, shape, and exposure time to enemies, repair scar rates also peaked to the south. Taken together, these results suggest that the expression of anti‐predatory traits along the geographic cline is governed by the interaction of two selective factors: temperature and predation, with the former acting as the ultimate control on the scope of adaptation both by escalating predation pressure in the southern part of B. carica's range and by physically limiting (to the north) and facilitating (to the south) the production of anti‐predatory traits. Feedbacks between temperature and predation thus causally interact to enable and drive, respectively, the observed geographic cline in energy‐intensive anti‐predatory shell traits.     

Representation of aquatic invertebrate communities in subfossil death assemblages sampled along a salinity gradient of western Uganda crater lakes

We analysed subfossil death assemblages of aquatic invertebrate communities in a salinity series of 35 western Uganda maar-crater lakes to evaluate their potential as biological indicators of past habitat conditions in paleo-environmental research. The study region encompasses the climatological and hydrological gradient between the dry floor and moist shoulders of the Edward-George branch of the East African Rift Valley, and includes mesotrophic to hyper-eutrophic, and shallow unstratified to deep meromictic lakes with a surface-water salinity range between 101 and 135 400 μS/cm. Focusing on non-chironomid aquatic invertebrates with good fossil preservation, we found that fossil larval remains of the Dipteran families Culicidae, Ephydridae, and Stratiomyidae are good indicators of saline environments. Our data further suggest that the abundances of Bryozoan statoblasts and Chaoboridae are indicative of, respectively, the fraction of the littoral zone covered by aquatic macrophytes and of lake trophic state, but a lake reference data set more specifically designed to cover variation in these environmental factors will be needed to determine the strength of these relationships. In these small, simple lake basins, recent death assemblages recovered from a single mid-lake surface-sediment sample provides a more complete inventory of local aquatic invertebrate communities and the distribution of species among lakes than exploratory live sampling of those taxa in a selection of littoral, benthic and pelagic habitats.     

Eutrophication in the Baltic Sea

Fish consumption is increasing globally. Overfishing puts pressure on fisheries, but aquaculture provides an alternative to satisfy the growing need for seafood. However, nutrient emissions from aquaculture contribute to eutrophication, and raising fish from the top of the food chain is inefficient. Here we use the approach of industrial ecology and report ImPACT decomposition analysis of the drivers of nutrient emissions to the Baltic Sea from rainbow trout aquaculture in Finland during 1980?2007. During this period, the nitrogen load studied increased markedly and was 522 tonnes in 2007. The phosphorus load quadrupled and then returned to its original level of about 65 tonnes. The Finnish population increased slightly, while the average affluence level increased significantly. Total salmonid consumption increased substantially during the period. The increasing percentage of imported salmonids and improvements in domestic aquaculture technology ended the period of strong growth of emissions in the 1980s. Decreasing the nutrient load through reductions in salmonid consumption in the future is unlikely, due to health benefits and consumer preferences. Replacing domestic production with import of salmonids raises questions regarding outsourcing of the environmental impact, and regarding rural development in Finland. Major improvements in production technology are not in sight. New perspectives on rainbow trout aquaculture may be needed, including using feed from the Baltic Sea, thus closing the nutrient cycle or changing consumption and production to herbivorous fish species.     

Methodological challenges in assessing the environmental status of a marine ecosystem: case study of the Baltic Sea

Assessments of the environmental status of marine ecosystems are increasingly needed to inform management decisions and regulate human pressures to meet the objectives of environmental policies. This paper addresses some generic methodological challenges and related uncertainties involved in marine ecosystem assessment, using the central Baltic Sea as a case study. The objectives of good environmental status of the Baltic Sea are largely focusing on biodiversity, eutrophication and hazardous substances. In this paper, we conduct comparative evaluations of the status of these three segments, by applying different methodological approaches. Our analyses indicate that the assessment results are sensitive to a selection of indicators for ecological quality objectives that are affected by a broad spectrum of human activities and natural processes (biodiversity), less so for objectives that are influenced by a relatively narrow array of drivers (eutrophications, hazardous substances). The choice of indicator aggregation rule appeared to be of essential importance for assessment results for all three segments, whereas the hierarchical structure of indicators had only a minor influence. Trend-based assessment was shown to be a useful supplement to reference-based evaluation, being independent of the problems related to defining reference values and indicator aggregation methodologies. Results of this study will help in setting priorities for future efforts to improve environmental assessments in the Baltic Sea and elsewhere, and to ensure the transparency of the assessment procedure.     

Evolution in a cline: the trilobite Triarthrus along an Ordovician depth gradient

Head proportions in the trilobite Triarthrus change by small but significant amounts with both lime and environment along a 'fossil' depth gradient in the Trenton Group of central New York. The putative zone species T. becki (Denmarkian Stage) and T. eatoni (post-Denmarkian; = T. becki , new synonymy) intermingle throughout the roughly two million year long sequence studied. The two represent extreme morphs in a graded spatio-temporal cline, not distinct species. Transgression as much as evolution accounts for their stratigraphic zonation. Clinal variation is expressed in the dependence of anatomical measures on the position of samples on a downslope transect parallel to bentonite beds, and on ordination score for community samples, a measure of environment that applies over time. Anatomy changes with environment in basically the same way it changes with time, and the amount of change over two million years in a given environment roughly equals the amount of change with environment at a given time. Compensated to a standard environment, anatomical measures follow a continuous trend that suggests an evolutionary random walk with non-random, directional drift. The case of Triarthrus illustrates that directional anatomical variation can have important components that are not strictly temporal and evolutionary in the normal sense.