Nutrient enrichment from large outlets of sewage and runoff from land is a major problem in many coastal areas. The study area, situated in the northern Baltic Proper just north of Stockholm on the Swedish coast, has received high amounts of phosphorus and nitrogen for over a century. The main part came from untreated sewage. In this study, we documented occurrence of attached brown perennial macroalgae Fucus vesiculosus thalli for geographical and depth distribution and compared this to older surveys from the same area. We also used herbarium material for comparing the cover and species composition of epibiota on F. vesiculosus. Nutrient reduction in the studied area began in the 1960’s. Today, phosphorus levels are similar to the 1930s. With reduced nutrient levels, Secchi depth improved from an average of 2.6 m in late 1960s to 4.5 m in the 2020 s. This increase in light penetration allowed for an increased depth distribution of F. vesiculosus. The nutrient reduction has also resulted in a natural recovery of F. vesiculosus in the area, and the innermost distribution is now back to where it was in the 1880s. The composition of epibiota on F. vesiculosus also showed changes, from a dense cover of filamentous algae to a high cover of filter-feeding invertebrates.

Genetic characteristics of populations can have substantial impacts on the adaptive potential of a species. Species are heterogeneous, often defined by variability at a range of scales including at the genetic, individual and population level. Using microsatellite genotyping, we characterize patterns underlying the genetic heterogeneity in marine macroalga Fucus vesiculosus, with a particular focus on two forms: attached and free-living. Here we demonstrate that sympatric populations representing the two forms display marked differences in characteristics of reproduction and genetic diversity. Asexual reproduction was ubiquitous in the free-living form despite being almost entirely absent in the attached form, while signals of polyploidy were common in both forms despite the distinct reproductive modes. Gene flow within and between the forms differed, with barriers to gene flow occurring between forms at various spatial scales due to the reproductive modes employed by individuals of each form. The divergent genetic characteristics of F. vesiculosus demonstrate that intraspecific differences can influence the properties of populations with consequential effects on the whole ecosystem. The differing genetic patterns and habitat requirements of the two forms define separate but closely associated ecological entities that will likely display divergent responses to future changes in environmental conditions. 

Sequencing of a mitochondrial intergenic spacer and 23S subunit was used to investigate the phylogeographic patterns in Fucus vesiculosus. Samples originated from 21 sites spanning six subbasins of the Baltic Sea. We identify a putative ancestral mitochondrial haplotype that entered the Baltic Sea from the Atlantic, colonising extensively throughout the species’ distribution. The dominance of this haplotype is seen in the low overall haplotype diversity (H_d = 0.29). Moreover, there is indication of few spatially aggregated patterns in the deeper demographic time scales (F_ct = 0.040; F_st = 0.049). Tajima’s D (−0.685, p-value 0.297) and Fu’s F_S (0.267, p-value 0.591) showed no significant signals of extreme demographic changes. The Baltic Sea free-living Fucus is confirmed as F. vesiculosus or a closely related species. Haplotype diversities are comparable between forms (attached H_d = 0.306; free-living H_d = 0.268). The relatively short temporal scale for colonisation alongside low variance in the Fucus mitochondrial genome results in a rather panmictic structure across the Baltic Sea. Our data suggest that the mitochondrial intergenic spacer and 23S poorly describe the evolutionary dynamics of Fucus spp. in such a young, postglacial environment, yet this concatenated-barcode advances our understanding of the colonisation dynamics of F. vesiculosus over deeper demographic timescales.

Ocean climate change strongly affects organisms and ecosystems, and the causes, consequences, and underlying mechanisms need to be documented. In the Baltic Sea, a marginal sea under severe eutrophication stress, a longer productive season, and changes in the phytoplankton community over the last few decades have likely impacted diet and condition of keystone species, from individual to population level. This study uses stable isotopes (δ¹³C, δ¹⁵N, and derived isotope niche metrics) to trace energy and nutrient flows in archived samples of blue mussel (Mytilus edulis trossulus) spanning 24 yr (1993–2016). We test if long-term changes in isotope and elemental composition in mussels, as well as population abundance and biomass, can be explained by changes in abiotic and biotic variables, using partial least square regressions and structural equation modeling. We found decreasing trends in δ13C and δ15N as well as in mean size and total biomass of mussels, but no unidirectional changes in their stoichiometry or condition index. Changes in isotope composition were best explained by nitrogen-fixing cyanobacteria, by increased terrestrial organic carbon from land runoff (reflecting precipitation) and by decreases in dissolved inorganic nitrogen (indicative of successful eutrophication mitigation) and in biomass of a mixotrophic ciliate species. The trophic niche (assessed from isotope niche) was included as the best predictor for both mussel body condition and the observed decline in their total biomass. This study reveals that altered trophic relationships from climate-induced changes in the productivity base may strongly impact keystone species, with potential knock-on effects on ecosystem functions. 

Handbokens syfte är att vara ett stöd inför planering och genomförande av projekt med avsikten att restaurera blåstångssamhällen i Östersjön. Alternativt, för att etablera ett blåstångssamhälle som en kompensationsåtgärd vid byggnationer och anläggningar i havet. En restaurering av ett blåstångssamhälle innebär att med insatta åtgärder hjälpa blåstång att återetablera sig i ett område där den tidigare funnits men försvunnit från på grund av mänsklig påverkan i form av till exempel övergödning och/eller föroreningar. En restaureringsåtgärd kan också innebära att den naturliga återhämtningen av ett blåstångssamhälle sker snabbare, dvs. att beståndets utbredning och/eller täthet ökar. Etablering av blåstångssamhällen som kompensationsåtgärd vid byggnationer i havet är högaktuellt i kustområden där exploateringstrycket är stort. Byggande av hamnar, vågbrytare och pirer skapar nya hårdbottnar som snabbt kan koloniseras av fintrådiga alger. Blåstången har sämre spridningsförmåga och kommer därmed att ha svårare att kolonisera nya ytor, vilket innebär att det kan ta lång tid innan ett tätt blåstångssamhälle etableras.

I handboken presenteras metoder för att etablera blåstångsamhällen i miljöer där den försvunnit, för att påskynda blåstångens naturliga kolonisering och etablering där den minskat, samt som kompensationsåtgärd vid byggnationer i havet. Inför planering av restaurerings- eller kompensationsåtgärder krävs grundläggande kunskaper om naturliga begränsningar och biologiska förutsättningar för blåstångens utbredning tillsammans med kunskap om blåstångens biologi och funktion i Östersjöns ekosystem. Denna bakgrundsinformation presenteras i första delen av handboken. Metodiken för att genomföra en restaureringsåtgärd av ett blåstångsamhälle är en process i flera steg och beskrivs i andra delen av handboken. Här beskrivs de förstudier av miljöförhållanden och biologiska förutsättningar som måste finnas innan en åtgärd att restaurera/nyetablera ett blåstångssamhälle initieras. Handboken bygger på sammanställda data från forskningslitteratur och miljöövervakning om blåstångens ekosystem i Östersjön samt författarnas egna erfarenheter från fältförsök, tester och observationer.

Handboken avser främst restaurering- och kompensationsåtgärder utifrån miljöförhållanden i Egentliga Östersjön, men bör kunna tillämpas i hela blåstångens utbredningsområde även i Bottenhavet med hänsyn till lokala förhållanden. Handboken är en handledning för att hjälpa utföraren att undvika kända problem vid återetablering av blåstång och ge konkreta förslag på metodik. Författarna vill dock påpeka att handboken inte garanterar att ett restaureringsförsök lyckas eftersom det fortfarande finns luckor i kunskapen om vad som krävs för att restaurera ett blåstångssamhälle. Arbetet med att genomföra fältstudier och underlag till handboken har finansierats av stiftelsen BalticSea2020 med stöd av Stockholms universitets Östersjöcentrum.

Flera äldre marinbotanister har skrivit om en frilevande form av blåstång. Nu arbetar forskare inom projektet FunkVeg på Stockholms och Helsingfors universitet med att kartlägga och dokumentera den frilevande blåstången som bildar viktiga livsmiljöer för många arter i Östersjöns kustnära vatten.

V i vill ta reda på vilken utbredning den frilevande formen av blåstång Fucus vesiculosus har i Sverige och Finland. Vår forskning strävar också efter att ta reda på vilken roll den spelar i Östersjöns grunda kustekosystem och hur bestånden förändras över tid och påverkas av klimatförändringen. Inventeringar i Tyskland på 2000-talet visade att de frilevande bestånden av blåstång gått tillbaka kraftigt, framförallt på grund av att övergödning och småskalig exploatering av grunda områden försämrat deras livsmiljö. Det ledde till att deras habitat klassats som hotat i Helsingforskommissionens rödlista för biotoper (Helcom 2013). Men vad vet vi egentligen om den frilevande blåstångens utbredning på en Östersjöskala och vilken funktion har den som habitat?

Fucus vesiculosus is common both on the tidal coasts of the North Atlantic and in the Baltic Sea, where it has adapted to low salinity and nontidal conditions over the last 7000 years. During the late 1970s and early 1980s, extensive declines of F. vesiculosus populations were reported in the Baltic Proper, mainly attributed to high nutrient loads. During the past 30-40 years, considerable efforts have been made to reduce nutrient runoff to coastal areas but few successful initiatives to restore F. vesiculosus populations have been performed. In this paper, we present how substratum manipulation, i.e. clean rocky surfaces, brushing rocks, Hildenbrandia rubra cover and different filamentous algae, as well as different algal exudates, affect the recruitment and survival of juvenile F.vesiculosus. Further, we show through a 5-year field experiment that it will take at least 4-5 years to reach reproductive age for F. vesiculosus in the Baltic Sea. We also present transplantation studies from two different areas, showing that epiphytic load, light, grazing and type of substratum are some of the factors that need to be taken into consideration in order to achieve successful restoration of F. vesiculosus.

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.

Fucus radicans is an endemic habitat-forming brown macroalga in the Baltic Sea that commonly complements its sexual reproduction with asexual reproduction. Asexual reproduction in F. radicans takes place through formation of adventitious branches (hereafter fragments), but the exact mechanisms behind it remain unknown.

We assessed experimentally the importance of two environmental factors determining the re-attachment success of F. radicans fragments. By combining different light conditions (daylength and irradiance; high or low light) and water temperature (+14°C and +4°C), we mimicked ambient light and temperature conditions of winter, spring/autumn and summer for F. radicans. Fragments were able to re-attach in all tested conditions. Temperature and light had an interactive impact on re-attachment: the combination of high temperature and high light level resulted in the highest re-attachment success, while light level had no effects on re-attachment success in cooler water temperature and the re-attachment success in high temperature under low light levels was very low.

The results suggest that rhizoid formation, and thus re-attachment success, may depend on the net primary production (metabolic balance) of the fragment. However, whether the re-attachment and asexual reproduction success simply depends on photosynthetic capacity warrants further mechanistic studies. Understanding the mechanisms of asexual reproduction in F. radicans is important in order to assess the dispersal capacity of this foundation species.

Many of the marine species that were introduced to the Baltic Sea during the Littorina stage (c. 8500-3000 years BP), e.g. Fucus vesiculosus and F. serratus, have adapted to the present low salinity. These marine species have gone from marine conditions into lower salinity environments. In this paper we ask why the recently discovered endemic brown alga Fucus radicans shows the opposite pattern. Fucus radicans is only present in the northern parts of the Baltic Sea, the low salinity Bothnian Sea (4-6 psu). Potentially, the fitness of F. radicans might be reduced in higher salinities if it is better adapted to brackish conditions. We hypothesize, however, that the southern distribution limit of F. radicans is set by biotic factors, e.g. competition with F. vesiculosus and higher grazing pressure by Idotea balthica and not by salinity. Our results show that the reproductive output of F. radicans is limited by low salinity (4 psu) but increases in higher salinities. However, the southern distribution limit, i.e. the northern Baltic Proper, is regulated by biotic factors, where the additive effects from shading by taller F. vesiculosus thalli and grazing on F. radicans by the isopod I. balthica limit the biomass production of F. radicans. We suggest that F. radicans still maintains marine traits due to its ability to propagate clonally and is restricted to the Bothnian Sea by interactions with F. vesiculosus and I. balthica. We also propose that increased precipitation due to climate change might affect the northern range limit and that the distribution of F. radicans could be expected to shift further south into the Baltic Proper.