Description: Pattern-triggered immunity (PTI) is an integral part of the innate immune system of many eukaryotic hosts, assisting in the defence against pathogen invasions. In plants and animals, PTI exerts a selective pressure on the microbiota that can alter community composition. However, the effect of PTI on the microbiota for non-model hosts, including seaweeds, remains unknown. Using quantitative polymerase chain reaction complemented with 16S rRNA gene and transcript amplicon sequencing, this study profiled the impact that PTI of the red seaweed Gracilaria gracilis has on its microbiota. PTI elicitation with agar oligosaccharides resulted in a significant reduction in the number of bacteria (by 〉75% within 72 h after treatment). However, the PTI elicitation did not cause any significant difference in the community diversity or structure. These findings demonstrated that PTI can be non-selective, and this might help to maintain a stable microbiota by uniformly reducing bacterial loads.

Description: Combined genetic, morphological and ontogenetic observations show that the circumarctic boreal green algal macrophyte Kornmannia leptoderma has expanded its distribution range into the Baltic Sea, on a German coastal section of 220 km length. The species is also again (or still) established at its former extreme southern distribution limit in the North Sea, the German island of Helgoland, where it has not been detected during the last four decades. Macroscopic visible sporophytes of K. leptoderma are nowadays present in the Baltic Sea and at Helgoland from February to September, while they were in the past only detected from February to May at Helgoland. This capacity for formation of sporophytes in summer correlates with the circumstance that K. leptoderma from the Baltic Sea can complete its life cycle at 15°C while several studies conducted decades ago with material from Helgoland and from Pacific coasts consistently reported an inhibition of the algal gametogenesis at temperatures that exceed 12°C. Possibly K. leptoderma has undergone adaptations that facilitate its spread into warmer environments, unless the Kornmannia present in the Baltic Sea and on Helgoland today represents a newly introduced cryptic species.

Description: Due to low salinity and lack of hard substrata, the Baltic Sea and Kattegat area and German and Danish North Sea coasts are characterized by a relatively low diversity of seaweeds. At the same time the areas are severely eutrophicated, which has caused extensive shifts in macroalgal communities toward opportunistic species. Unattached seaweed communities dominated by Furcellaria lumbricalis, which have been a resource for hydrocolloid production since the 1940s, have been severely reduced due to eutrophication and unsustainable harvesting and are nowadays only exploited commercially in Estonia. On the other hand, the biomass of opportunistic seaweeds of various red, green and brown algal genera has increased. They cause ecological problems, are a nuisance on many tourist beaches and constitute at the same time a potential bioresource that is so far only exploited to a limited extent for production of energy and fertilizer. Commercial seaweed cultivation is largely focused on Saccharina latissima and still very limited, but is currently being expanded as a compensation measure for sea-based fish aquaculture. Also land-based seaweed cultivation is primarily employed for recycling of nutrients in tank animal aquaculture, but in most cases so far only on an experimental scale.

Description: Seaweeds are colonized by a microbial community, which can be directly linked to their performance. This community is shaped by an interplay of stochastic and deterministic processes, including mechanisms which the holobiont host deploys to manipulate its associated microbiota. The Anna Karenina principle predicts that when a holobiont is exposed to suboptimal or stressful conditions, these host mechanisms may be compromised. This leads to a relative increase of stochastic processes that may potentially result in the succession of a microbial community harmful to the host. Based on this principle, we used the variability in microbial communities (i.e., beta diversity) as a proxy for stability within the invasive holobiont Gracilaria vermiculophylla during a simulated invasion in a common garden experiment. Independent of host range, host performance declined at elevated temperature (22°C) and disease incidence and beta diversity increased. Under thermally stressful conditions, beta diversity increased more in epibiota from native populations, suggesting that epibiota from non-native holobionts are thermally more stable. This pattern reflects an increase in deterministic processes acting on epibiota associated with non-native hosts, which in the setting of a common garden can be assumed to originate from the host itself. Therefore, these experimental data suggest that the invasion process may have selected for hosts better able to maintain stable microbiota during stress. Future studies are needed to identify the underlying host mechanisms.

Description: Supralittoral and shallow water seaweed communities are particularly exposed to impacts such as climate change and disturbance by humans. Therefore, their classification, the study of composition, and the monitoring of their structural changes are particularly important. A phytosociological survey of the supralittoral and upper sublittoral vegetation of the South West Baltic Sea revealed eight phytobenthos communities with two variants comprising 35 taxa of macrophytes (18 taxa of Chlorophyta, 13 taxa of Rhodophyta and four taxa of Phaeophyceae, Ochrophyta). Five of the eight communities were dominated by Ulvales (Ulva intestinalis, Kornmannia leptoderma, and three Blidingia species), the other three by Fucus vesiculosus. Most Fucus vesiculosus-dominated communities contained U. intestinalis and U. linza as subdominants. Only one of the communities had until now been described as an association ( Ulvetum intestinalis Feldman 1937). The syntaxonomic composition of the investigated vegetation includes both phytocenoses with the domination of green algae ( Ulvetum intestinalis Feldman 1937 and communities of Blidingia marginata, unidentified Blidingia spp. and Kornmannia leptoderma), as well as a number of communities dominated by Fucus vesiculosus. Mainly boreal Atlantic species and cosmopolitans make up the bulk of the species in these associations.

Description: 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.

Description: The Kiel Canal is one of the world’s most frequently used inland waterways and connects the SW Baltic Sea with the Wadden Sea. At the same time, the canal is a highly eutrophicated environment that is characterized by salinities that range from 3 to 16. This brackish character could make the Kiel Canal an important stepping stone for the introductions of species into the inner Baltic Sea. It could also hinder the identification of native and introduced species, given the fact that salinity sometimes severely affects algal morphology. Here we report on a survey of introduced and native seaweed species in the canal, focusing on the dominant groups, which are Fucales and Ulvales. Of the Fucales, the introduced species Fucus evanescens was detected nearly exclusively inside the canal, while Fucus vesiculosus dominated rockweed communities directly outside the sluice gates. Morphological analysis and genetic barcoding distinguished three species of Ulvales, Ulva linza, Ulva intestinalis and an unknown and possibly introduced species of the genus Blidingia. Species distributions and – in the case of U. intestinalis – branching patterns were clearly affected by salinity, while thallus sizes appeared to be affected by the specific eutrophication status of sites within the canal.

Description: Communities are shaped by scale dependent processes. To study the diversity and variation of microbial communities across scales, the invasive and widespread seaweed Agarophyton vermiculophyllum presents a unique opportunity. We characterized pro‐ and eukaryotic communities associated with this holobiont across its known distribution range, which stretches over the northern hemisphere. Our data reveal that community composition and diversity in the holobiont vary at local but also larger geographic scales. While processes acting at the local scale (i.e., within population) are the main structuring drivers of associated microbial communities, changes in community composition also depend on processes acting at larger geographic scales. Interestingly, the largest analysed scale (i.e., native and non‐native ranges) explained variation in the prevalence of predicted functional groups, which could suggest a functional shift in microbiota occurred over the course of the invasion process. While high variability in microbiota at the local scale supports A. vermiculophyllum to be a generalist host, we also identified a number of core taxa. These geographically independent holobiont members imply that cointroduction of specific microbiota may have additionally promoted the invasion process.

Description: Rapid evolution of non-native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction and magnitude of phenotypic evolution during invasion has been underestimated. We explored the utility of niche-shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe and northwestern Africa within the last 100 years. A genetically-informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non-native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non-native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field-collected and 325 common-garden thalli from 40 locations and as predicted, non-native populations had greater tolerance for ecologically-relevant extreme heat (40°C) than did Japanese source populations. Non-native populations also had greater tolerance for cold and low-salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: populations from warmer, lower-latitude estuaries had greater heat tolerance than did populations from colder, higher-latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non-native marine species. Genetically-informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions, and may help to resolve debate over the frequency of niche conservatism versus rapid adaptation during invasion.

Description: 1. Epimicrobial communities on seaweed surfaces usually contain not only potentially pathogenic but also potentially beneficial micro‐organisms. Capacity of terrestrial plants for chemically mediated recruitment, that is, “gardening” of bacterial communities in the rhizosphere was recently demonstrated. Empirical evidence directly linking such chemical “gardening” with the beneficial role of gardened microbes in terrestrial plants is rare and largely missing for aquatic macrophytes. 2. Here, we demonstrate that our model invasive seaweed holobiont Agarophyton vermiculophyllum possesses beneficial microbiota on its surface that provide protection from bacterial pathogens. Metabolites from the algal holobiont’s surface reduced settlement of opportunistic pathogens but attracted protective epibacterial settlement. 3. We tested 58 different bacterial species (isolated from the surface of A. vermiculophyllum) individually in tip bleaching assays. Kordia algicida was identified as a “significant pathogen” inducing a bleaching disease. In addition, nine other species significantly reduced the risk of algal bleaching and were thus “significantly protective”. Additionally, two “potential pathogens” and 10 “potential protectors” were identified. When 19 significant and potential protectors and 3 significant and potential pathogens were tested together, the protective strains fully prevented bleaching, suggesting that a component of A. vermiculophyllum’s epimicrobiome provides an associational defence against pathogens. Chemically mediated selective recruitment of microbes was demonstrated in bioassays, where A. vermiculophyllum surface metabolites attracted the settlement of protective strains, but reduced settlement of pathogens. 4. Synthesis. The capacity of an aquatic macrophyte to chemically “garden” protective micro‐organisms to the benefit of strengthened disease resistance is demonstrated for the first time. Such a role of surface chemistry in “gardening” of microbes as found in the current study could also be applicable to other host plant—microbe interactions. Our results may open new avenues towards manipulation of the surface microbiome of seaweeds via chemical “gardening,” enhancing sustainable production of healthy seaweeds.