Kelp forest community structure and demography in Kongsfjorden (Svalbard) across 25 years of Arctic warming.

The Arctic archipelago of Svalbard is a hotspot of global warming and many fjords experience a continuous increase in seawater temperature and glacial melt while sea-ice cover declines. In 1996/1998, 2012-2014, and 2021 macroalgal biomass and species diversity were quantified at the study site Hansneset, Kongsfjorden (W-Spitsbergen) in order to identify potential changes over time. In 2021, we repeated the earlier studies by stratified random sampling (1 × 1 m2, n = 3) along a sublittoral depth transect (0, 2.5, 5, 10, and 15 m) and investigated the lower depth limits of dominant brown algae between 3 and 19 m. The maximum fresh weight (FW) of all seaweeds was 11.5 kg m-2 at 2.5 m and to 99.9% constituted of kelp. Although biomass distribution along the depth transect in 2021 was not significantly different compared to 2012/2013, the digitate kelp community (Laminaria digitata/Hedophyllum nigripes) had transformed into an Alaria esculenta-dominated kelp forest. Consequently, a pronounced shift in kelp forest structure occurred over time as we demonstrate that biomass allocation to thallus parts is kelp species-specific. Over the past decade, kelp demography changed and in 2021 a balanced age structure of kelps (juveniles plus many older kelp individuals) was only apparent at 2.5 m. In addition, the abundances and lower depth limits of all dominant brown algae declined noticeably over the last 25 years while the red algal flora abundance remained unchanged at depth. We propose that the major factor driving the observed changes in the macroalgal community are alterations in underwater light climate, as in situ data showed increasing turbidity and decreasing irradiance since 2012 and 2017, respectively. As a consequence, the interplay between kelp forest retreat to lower depth levels caused by coastal darkening and potential macroalgal biomass gain with increasing temperatures will possibly intensify in the future with unforeseen consequences for melting Arctic coasts and fjord ecosystem services.

A conceptual framework to help choose appropriate blue nature-based solutions.

Biodiversity loss and climate change have severely impacted ecosystems and livelihoods worldwide, compromising access to food and water, increasing disaster risk, and affecting human health globally. Nature-based Solutions (NbS) have gained interest in addressing these global societal challenges. Although much effort has been directed to NbS in urban and terrestrial environments, the implementation of NbS in marine and coastal environments (blue NbS) lags. The lack of a framework to guide decision-makers and practitioners through the initial planning stages appears to be one of the main obstacles to the slow implementation of blue NbS. To address this, we propose an integrated conceptual framework, built from expert knowledge, to inform the selection of the most appropriate blue NbS based on desired intervention objectives and social-ecological context. Our conceptual framework follows a four incremental steps structure: Step 1 aims to identify the societal challenge(s) to address; Step 2 highlights ecosystem services and the underlying biodiversity and ecological functions that could contribute to confronting the societal challenge(s); Step 3 identify the specific environmental context the intervention needs to be set within (e.g. the spatial scale the intervention will operate within, the ecosystem's vulnerability to stressors, and its ecological condition); and Step 4 provides a selection of potential blue NbS interventions that would help address the targeted societal challenge(s) considering the context defined through Step 3. Designed to maintain, enhance, recover, rehabilitate, or create ecosystem services by supporting biodiversity, the blue NbS intervention portfolio includes marine protection (i.e., fully, highly, lightly, and minimally protected areas), restorative activities (i.e., active, passive, and partial restoration; rehabilitation of ecological function and ecosystem creation), and other management measures (i.e., implementation and enforcement of regulation). Ultimately, our conceptual framework guides decision-makers toward a versatile portfolio of interventions that cater to the specific needs of each ecosystem rather than imposing a rigid, one-size-fits-all model. In the future, this framework needs to integrate socio-economic considerations more comprehensively and be kept up-to-date by including the latest scientific information.

18S rDNA gene metabarcoding of microeukaryotes and epi-endophytes in the holobiome of seven species of large brown algae.

Brown algae (Phaeophyceae) are habitat-forming species in coastal ecosystems and include kelp forests and seaweed beds that support a wide diversity of marine life. Host-associated microbial communities are an integral part of phaeophyte biology, and whereas the bacterial microbial partners have received considerable attention, the microbial eukaryotes associated with brown algae have hardly been studied. Here, we used broadly targeted "pan-eukaryotic" primers (metabarcoding) to investigate brown algal-associated eukaryotes (the eukaryome). Using this approach, we aimed to investigate the eukaryome of seven large brown algae that are important and common species in coastal ecosystems. We also aimed to assess whether these macroalgae harbor novel eukaryotic diversity and to ascribe putative functional roles to the host-associated eukaryome based on taxonomic affiliation and phylogenetic placement. We detected a significant diversity of microeukaryotic and algal lineages associated with the brown algal species investigated. The operational taxonomic units (OTUs) were taxonomically assigned to 10 of the eukaryotic major supergroups, including taxonomic groups known to be associated with seaweeds as epibionts, endobionts, parasites, and commensals. Additionally, we revealed previously unrecorded sequence types, including novel phaeophyte OTUs, particularly in the Fucus spp. samples, that may represent fucoid genomic variants, sequencing artifacts, or undescribed epi-/endophytes. Our results provide baseline data and technical insights that will be useful for more comprehensive seaweed eukaryome studies investigating the evidently lineage-rich and functionally diverse symbionts of brown algae.

Effects of depth and overgrowth of ephemeral macroalgae on a remote subtidal NE Atlantic eelgrass (Zostera marina) community.

We conducted a short-term field sampling complemented with time integrating stable isotope analysis to holistically investigate status and ecological interactions in a remote NE Atlantic Zostera marina meadow. We found high nutrient water concentrations, large biomass of fast-growing, ephemeral macroalgae, low abundance, and biodiversity of epifauna and a food web with thornback ray (Raja clavata) as intermediate and cod (Gadus morhua) as top predator. We observed no variation with increasing depth (3.5-11 m) except for decreasing shoot density and biomass of Zostera and macroalgae. Our results indicate that the Finnøya Zostera ecosystem is eutrophicated. During the past three to four decades, nutrients from aquaculture have steadily increased to reach 75% of anthropogenic input while the coastal top predator cod has decreased by 50%. We conclude that bottom-up regulation is a predominant driver of change since top-down regulation is generally weak in low density and exposed Zostera ecosystems such as Finnøya.

The Saccharina latissima microbiome: Effects of region, season, and physiology.

Introduction: Saccharina latissima is a canopy-forming species of brown algae and, as such, is considered an ecosystem engineer. Several populations of this alga are exploited worldwide, and a decrease in the abundance of S. latissima at its southern distributional range limits has been observed. Despite its economic and ecological interest, only a few data are available on the composition of microbiota associated with S. latissima and its role in algal physiologyn. Methods: We studied the whole bacterial community composition associated with S. latissima samples from three locations (Brittany, Helgoland, and Skagerrak) by 16S metabarcoding analyses at different scales: algal blade part, regions, season (at one site), and algal physiologic state. Results and Discussion: We have shown that the difference in bacterial composition is driven by factors of decreasing importance: (i) the algal tissues (apex/meristem), (ii) the geographical area, (iii) the seasons (at the Roscoff site), and (iv) the algal host's condition (healthy vs. symptoms). Overall, Alphaproteobacteria, Gammaproteobacteria, and Bacteroidia dominated the general bacterial communities. Almost all individuals hosted bacteria of the genus Granulosicoccus, accounting for 12% of the total sequences, and eight additional core genera were identified. Our results also highlight a microbial signature characteristic for algae in poor health independent of the disease symptoms. Thus, our study provides a comprehensive overview of the S. latissima microbiome, forming a basis for understanding holobiont functioning.

Homogenization and miniaturization of habitat structure in temperate marine forests.

Humans are rapidly transforming the structural configuration of the planet's ecosystems, but these changes and their ecological consequences remain poorly quantified in underwater habitats. Here, we show that the loss of forest-forming seaweeds and the rise of ground-covering 'turfs' across four continents consistently resulted in the miniaturization of underwater habitat structure, with seascapes converging towards flattened habitats with smaller habitable spaces. Globally, turf seascapes occupied a smaller architectural trait space and were structurally more similar across regions than marine forests, evidencing habitat homogenization. Surprisingly, such habitat convergence occurred despite turf seascapes consisting of vastly different species richness and with different taxa providing habitat architecture, as well as across disparate drivers of marine forest decline. Turf seascapes contained high sediment loads, with the miniaturization of habitat across 100s of km in mid-Western Australia resulting in reefs retaining an additional ~242 million tons of sediment (four orders of magnitude more than the sediments delivered fluvially annually). Together, this work demonstrates that the replacement of marine forests by turfs is a generalizable phenomenon that has profound consequences for the ecology of temperate reefs.

DNA barcoding and mucilage ducts in the stipe reveal the presence of Hedophyllum nigripes (Laminariales, Phaeophyceae) in Kongsfjorden (Spitsbergen).

The Arctic Ocean is a unique ecosystem hosting a biodiversity that has not yet been elucidated in full detail. There is increasing evidence that there are more kelp species constricted to Arctic/sub-Arctic habitats hitherto not well investigated, such as Hedophyllum nigripes, which is morphologically very similar to cold-temperate Laminaria digitata. Hedophyllum nigripes was originally described as L. nigripes by Agardh from Spitsbergen but has often been misidentified as L. digitata in the European Arctic. We initiated a systematic algal survey along a depth gradient (0-7.5 m) in Kongsfjorden (Spitsbergen) in June and July 2015 and thereby confirmed a predominant presence of H. nigripes (73%). Hedophyllum nigripes is occurring between 0 and 7.5 m while L. digitata was most abundant at 2.5 m depth. Hedophyllum nigripes individuals were generally younger (2.3 vs. 3.6 years) and stipe and blade length shorter (31 vs. 54 cm and 76 vs. 96 cm, respectively) compared to L. digitata. A combination of molecular (COI-5P) and morpho-anatomical tools (presence of sori and mucilage ducts in the stipe) was used to differentiate specimens of H. nigripes and L. digitata. Both kelp species were indistinguishable in most cases by external blade and stipe morphology. The different blade shapes represented different ontogenetic stages rather than phenotypic plasticity. The presence of mucilage ducts in the stipe was correlated with H. nigripes and changed with depth from 17%, 36%, and 85% at 2.5, 5, and 7.5 m, respectively. In addition, all summer fertile specimens were L. digitata.

Green gravel: a novel restoration tool to combat kelp forest decline.

Kelp forests are in decline globally and large-scale intervention could be required to halt the loss of these valuable ecosystems. To date kelp forest restoration has had limited success and been expensive and unable to address the increasing scale of ecosystem deterioration. Here we developed and tested a new approach: "green gravel". Small rocks were seeded with kelp and reared in the laboratory until 2-3 cm, before out-planting to the field. The out-planted kelp had high survival and growth over 9 months, even when dropped from the surface. This technique is cheap, simple, and does not require scuba diving or highly trained field workers. It can be up-scaled to treat large areas or even used to introduce genes from more resilient kelp populations onto vulnerable reefs. Green gravel thus overcomes some of the current major limitations of kelp restoration and provides a promising new defense against kelp forest decline.

Carbon export is facilitated by sea urchins transforming kelp detritus.

With the increasing imperative for societies to act to curb climate change by increasing carbon stores and sinks, it has become critical to understand how organic carbon is produced, released, transformed, transported, and sequestered within and across ecosystems. In freshwater and open-ocean systems, shredders play a significant and well-known role in transforming and mobilizing carbon, but their role in the carbon cycle of coastal ecosystems is largely unknown. Marine plants such as kelps produce vast amounts of detritus, which can be captured and consumed by shedders as it traverses the seafloor. We measured capture and consumption rates of kelp detritus by sea urchins across four sampling periods and over a range of kelp detritus production rates and sea urchin densities, in northern Norway. When sea urchin densities exceeded 4 m-2, the sea urchins captured and consumed a high percentage (ca. 80%) of kelp detritus on shallow reefs. We calculated that between 1.3 and 10.8 kg of kelp m-2 are shredded annually from these reefs. We used a hydrodynamic dispersal model to show that transformation of kelp blades to sea urchin feces increased its export distance fourfold. Our findings show that sea urchins can accelerate and extend the export of carbon to neighboring areas. This collector-shredder pathway could represent a significant flow of small particulate carbon from kelp forests to deep-sea areas, where it can subsidize benthic communities or contribute to the global carbon sink.

Detrital carbon production and export in high latitude kelp forests.

The production and fate of seaweed detritus is a major unknown in the global C-budget. Knowing the quantity of detritus produced, the form it takes (size) and its timing of delivery are key to understanding its role as a resource subsidy to secondary production and/or its potential contribution to C-sequestration. We quantified the production and release of detritus from 10 Laminaria hyperborea sites in northern Norway (69.6° N). Kelp biomass averaged 770 ± 100 g C m-2 while net production reached 499 ± 50 g C m-2 year-1, with most taking place in spring when new blades were formed. Production of biomass was balanced by a similar formation of detritus (478 ± 41 g C m-2 year-1), and both were unrelated to wave exposure when compared across sites. Distal blade erosion accounted for 23% of the total detritus production and was highest during autumn and winter, while dislodgment of whole individuals and/or whole blades corresponded to 24% of the detritus production. Detachment of old blades constituted the largest source of kelp detritus, accounting for > 50% of the total detrital production. Almost 80% of the detritus from L. hyperborea was thus in the form of whole plants or blades and > 60% of that was delivered as a large pulse within 1-2 months in spring. The discrete nature of the delivery suggests that the detritus cannot be retained and consumed locally and that some is exported to adjacent deep areas where it may subsidize secondary production or become buried into deep marine sediments as blue carbon.

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.

Efficient coralline algal psbA mini barcoding and High Resolution Melt (HRM) analysis using a simple custom DNA preparation.

Coralline algae form extensive maerl and rhodolith habitats that support a rich biodiversity. Calcium carbonate harvesting as well as trawling activities threatens this ecosystem. Eleven species were recorded so far as maerl-forming in NE Atlantic, but identification based on morphological characters is unreliable. As for most red algae, we now use molecular characters to resolve identification of these taxa. However, obtaining DNA sequences requires time and resource demanding methods. The purpose of our study was to improve methods for achieving simple DNA extraction, amplification, sequencing and sequence analysis to allow robust identification of maerl species and other coralline algae. Our novel and easy DNA preparation method for coralline algae was of sufficient quality for qPCR amplification and sequencing of all 47 tested samples. The new psbA qPCR assay successfully amplified a 350 bp fragment identifying six species and uncovering two new Operational Taxonomic Units. Molecular results were corroborated with anatomical examination using i.e. scanning electron microscopy. Finally, the qPCR assay was coupled with High Resolution Melt analysis that successfully differentiated the closely related species Lithothamnion erinaceum and L. cf. glaciale. This DNA preparation and qPCR technique should vitalize coralline research by reducing time and cost associated with molecular systematics.

A snap shot of the short-term response of crustaceans to macrophyte detritus in the deep Oslofjord.

A test deployment of a time-lapse camera lander in the deep Oslofjord (431 m) was used to obtain initial information on the response of benthic fauna to macroalgal debris. Three macroalgal species were used on the lander baited plate: Fucus serratus, Saccharina latissima and Laminaria hyperborea and observed during 41.5 hours. The deep-water shrimp Pandalus borealis were attracted to the macroalgae rapidly (3 min after the lander reached the seafloor), followed by amphipods. Shrimp abundances were significantly higher in areas covered by macroalgae compared to the adjacent seafloor and the number of shrimp visiting the macroalgae increased with time. Amphipods arrived 13 hours later and were observed mainly on decaying L. hyperborea. The abundance of amphipods on L. hyperborea increased rapidly, reaching a peak at 31 h after deployment. These initial observations suggest that debris from kelp forests and other macroalgal beds may play an important role in fuelling deep benthic communities in the outer Oslofjord and, potentially, enhance secondary production of commercial species such as P. borealis.

Biodiversity mediates top-down control in eelgrass ecosystems: a global comparative-experimental approach.

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Distribution, structure and function of Nordic eelgrass (Zostera marina) ecosystems: implications for coastal management and conservation.

This paper focuses on the marine foundation eelgrass species, Zostera marina, along a gradient from the northern Baltic Sea to the north-east Atlantic. This vast region supports a minimum of 1480 km2 eelgrass (maximum >2100 km2), which corresponds to more than four times the previously quantified area of eelgrass in Western Europe.Eelgrass meadows in the low salinity Baltic Sea support the highest diversity (4-6 spp.) of angiosperms overall, but eelgrass productivity is low (<2 g dw m-2 d-1) and meadows are isolated and genetically impoverished. Higher salinity areas support monospecific meadows, with higher productivity (3-10 g dw m-2 d-1) and greater genetic connectivity. The salinity gradient further imposes functional differences in biodiversity and food webs, in particular a decline in number, but increase in biomass of mesograzers in the Baltic.Significant declines in eelgrass depth limits and areal cover are documented, particularly in regions experiencing high human pressure. The failure of eelgrass to re-establish itself in affected areas, despite nutrient reductions and improved water quality, signals complex recovery trajectories and calls for much greater conservation effort to protect existing meadows.The knowledge base for Nordic eelgrass meadows is broad and sufficient to establish monitoring objectives across nine national borders. Nevertheless, ensuring awareness of their vulnerability remains challenging. Given the areal extent of Nordic eelgrass systems and the ecosystem services they provide, it is crucial to further develop incentives for protecting them. © 2014 The Authors. Aquatic Conservation: Marine and Freshwater Ecosystems published by John Wiley & Sons, Ltd.

Seaweeds in cold seas: evolution and carbon acquisition.

Much evidence suggests that life originated in hydrothermal habitats, and for much of the time since the origin of cyanobacteria (at least 2.5 Ga ago) and of eukaryotic algae (at least 2.1 Ga ago) the average sea surface and land surface temperatures were higher than they are today. However, there have been at least four significant glacial episodes prior to the Pleistocene glaciations. Two of these (approx. 2.1 and 0.7 Ga ago) may have involved a 'Snowball Earth' with a very great impact on the algae (sensu lato) of the time (cyanobacteria, Chlorophyta and Rhodophyta) and especially those that were adapted to warm habitats. By contrast, it is possible that heterokont, dinophyte and haptophyte phototrophs only evolved after the Carboniferous-Permian ice age (approx. 250 Ma ago) and so did not encounter low (

Mechanistic interpretation of carbon isotope discrimination by marine macroalgae and seagrasses.

The literature, and previously unpublished data from the authors' laboratories, shows that the δ13C of organic matter in marine macroalgae and seagrasses collected from the natural environment ranges from -3 to -35‰. While some marine macroalgae have δ13C values ranging over more than 10‰ within the thallus of an individual (some brown macroalgae), in other cases the range within a species collected over a very wide geographical range is only 5‰ (e.g. the red alga Plocamium cartilagineum which has values between -30 and -35‰). The organisms with very negative δ13C (lower than -30‰) are mainly subtidal red algae, with some intertidal red algae and a few green algae; those with very positive δ13C values (higher than -10‰) are mainly green macroalgae and seagrasses, with some red and brown macroalgae. The δ13C value correlates primarily with taxonomy and secondarily with ecology. None of the organisms with δ13C values lower than -30‰ have pyrenoids. Previous work showed a good correlation between δ13C values lower than -30‰ and the lack of CO2 concentrating mechanisms for several species of marine red algae. The extent to which the low δ13C values are confined to organisms with diffusive CO2 entry is discussed. Diffusive CO2 entry could also occur in organisms with higher δ13C values if diffusive conductance was relatively low. The photosynthesis of organisms with δ13C values more positive than -10‰ (i.e. more positive than the δ13C of CO2 in seawater) must involve HCO3- use.