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  • 1. Angove, Charlotte
    et al.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Gustafsson, Camilla
    Assessing the efficiencies and challenges for nutrient uptake by aquatic plants2018In: Journal of Experimental Marine Biology and Ecology, ISSN 0022-0981, E-ISSN 1879-1697, Vol. 507, p. 23-30Article in journal (Refereed)
    Abstract [en]

    Aquatic plant meadows are valuable components to the 'coastal filter' and it is important to understand the processes that drive their ability to cycle nutrients. However, at present, the field-based evidence for understanding the drivers of nutrient uptake by plants is lacking. This study aimed to investigate how well individual shoots of aquatic plants could meet their nitrogen demands using the sediment nutrient pool (porewater ammonium) and to explore which traits helped to facilitate such uptake. Several species were investigated in shallow, submerged (2-4 m) mixed-species communities in the northern Baltic Sea using incubation experiments with enriched ammonium. After a 3.5 h incubation time, individuals were collected and analysed for nitrogen (% DW) and N-15 (at-%) concentrations. Uptake by plants was calculated per unit nitrogen in response to the N-15 labelled source and to overall nitrogen availability. Background porewater ammonium availability was highly variable between individual plants. Species identity did not significantly affect uptake metrics and the effect of ambient porewater availability was weak. As biomass increased there were significant logarithmic declines in the 95th quantiles of nutrient uptake rates, ambient porewater nutrient availability and aboveground nitrogen tissue concentrations (% DW). Such findings suggested that uptake rates of plants were significantly demand driven and the nutrient conditions of the porewater were significantly driven by the demands of the plant. Findings parameterised the unfulfilled potential for some aquatic plants to cycle nutrients more efficiently and highlighted the potential importance of access to new nutrient sources as a way of enhancing nutrient cycling by aquatic plants. Plant traits and community properties such as the activity of infauna could facilitate such an access and are likely important for nutrient uptake.

  • 2. Attard, K. M.
    et al.
    Rodil, Iván F.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Berg, P.
    Norkko, J.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Glud, R. N.
    Seasonal metabolism and carbon export potential of a key coastal habitat: The perennial canopy-forming macroalga Fucus vesiculosus2019In: Limnology and Oceanography, ISSN 0024-3590, E-ISSN 1939-5590, Vol. 64, no 1, p. 149-164Article in journal (Refereed)
    Abstract [en]

    The important role of macroalgal canopies in the oceanic carbon (C) cycle is increasingly being recognized, but direct assessments of community productivity remain scarce. We conducted a seasonal study on a sublittoral Baltic Sea canopy of the brown alga Fucus vesiculosus, a prominent species in temperate and Arctic waters. We investigated community production on hourly, daily, and seasonal timescales. Aquatic eddy covariance (AEC) oxygen flux measurements integrated similar to 40 m(2) of the seabed surface area and documented considerable oxygen production by the canopy year-round. High net oxygen production rates of up to 35 +/- 9 mmol m(-2) h(-1) were measured under peak irradiance of similar to 1200 mu mol photosynthetically active radiation (PAR) m(-2) s(-1) in summer. However, high rates > 15 mmol m(-2) h(-1) were also measured in late winter (March) under low light intensities < 250 mu mol PAR m(-2) s(-1) and water temperatures of similar to 1 degrees C. In some cases, hourly AEC fluxes documented an apparent release of oxygen by the canopy under dark conditions, which may be due to gas storage dynamics within internal air spaces of F. vesiculosus. Daily net ecosystem metabolism (NEM) was positive (net autotrophic) in all but one of the five measurement campaigns (December). A simple regression model predicted a net autotrophic canopy for two-thirds of the year, and annual canopy NEM amounted to 25 mol O-2 m(-2) yr(-1), approximately six-fold higher than net phytoplankton production. Canopy C export was similar to 0.3 kg C m(-2) yr(-1), comparable to canopy standing biomass in summer. Macroalgal canopies thus represent regions of intensified C assimilation and export in coastal waters.

  • 3. Attard, Karl M.
    et al.
    Rodil, Iván F.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Glud, Ronnie N.
    Berg, Peter
    Norkko, Joanna
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Seasonal ecosystem metabolism across shallow benthic habitats measured by aquatic eddy covariance2019In: Limnology and oceanography letters, ISSN 2378-2242, Vol. 4, no 3, p. 79-86Article in journal (Refereed)
    Abstract [en]

    Shallow benthic habitats are hotspots for carbon cycling and energy flow, but metabolism (primary production and respiration) dynamics and habitat-specific differences remain poorly understood. We investigated daily, seasonal, and annual metabolism in six key benthic habitats in the Baltic Sea using similar to 2900h of in situ aquatic eddy covariance oxygen flux measurements. Rocky substrates had the highest metabolism rates. Habitat-specific annual primary production per m(2) was in the order Fucus vesiculosus canopy>Mytilus trossulus reef>Zostera marina canopy>mixed macrophytes canopy>sands, whereas respiration was in the order M. trossulus>F. vesiculosus>Z. marina>mixed macrophytes> sands>aphotic sediments. Winter metabolism contributed 22-31% of annual rates. Spatial upscaling revealed that benthic habitats drive >90% of ecosystem metabolism in waters <= 5 m depth, highlighting their central role in carbon and nutrient cycling in shallow waters.

  • 4. Cummings, Vonda J.
    et al.
    Hewitt, Judi E.
    Thrush, Simon F.
    Marriott, Peter M.
    Halliday, N. Jane
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Linking Ross Sea Coastal Benthic Communities to Environmental Conditions: Documenting Baselines in a Spatially Variable and Changing World2018In: Frontiers in Marine Science, E-ISSN 2296-7745, Vol. 5, article id 232Article in journal (Refereed)
    Abstract [en]

    Understanding the functionality of marine benthic ecosystems, and how they are influenced by their physical environment, is fundamental to realistically predicting effects of future environmental change. The Antarctic faces multiple environmental pressures associated with greenhouse gas emissions, emphasizing the need for baseline information on biodiversity and the bio-physical processes that influence biodiversity. We describe a survey of shallow water benthic communities at eight Ross Sea locations with a range of environmental characteristics. Our analyses link coastal benthic community composition to seafloor habitat and sedimentary parameters and broader scale features, at locations encompassing considerable spatial extent and variation in environmental characteristics (e.g., seafloor habitat, sea ice conditions, hydrodynamic regime, light). Our aims were to: (i) document existing benthic communities, habitats and environmental conditions against which to assess future change, (ii) investigate the relationships between environmental and habitat characteristics and benthic community structure and function, and (iii) determine whether these relationships were dependent on spatial extent. A very high percentage (>95%) of the between-location variability in macro- or epifaunal community composition was explainable using multi-scale environmental variables. The explanatory power varied depending on the scale of influence of the environmental variables measured (fine and medium-scale habitat, broad scale), and with community type. However, the inclusion of parameters at all scales produced the most powerful model for both communities. Ice duration, ice thickness and snow cover were important broad scale variables identified that directly relate to climate change. Even when using only habitat-scale variables, extending the spatial scale of the study from three locations covering 32 km to eight locations covering ~340 km increased the degree of explanatory power from 18–32 to 64–78%. The increase in explanatory power with spatial extent lends weight to the possibility of using an indirect “space for time” substitution approach for future predictions of the effects of change on these coastal marine ecosystems. Given the multiple and interacting drivers of change in Antarctic coastal ecosystems a multidisciplinary, long term, repeated observation approach will be vital to both improve and test predictions of how coastal communities will respond to environmental change.

  • 5.
    Ehrnsten, Eva
    et al.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Timmermann, Karen
    Gustafsson, Bo G.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Benthic-pelagic coupling in coastal seas - Modelling macrofaunal biomass and carbon processing in response to organic matter supply2019In: Journal of Marine Systems, ISSN 0924-7963, E-ISSN 1879-1573, Vol. 196, p. 36-47Article in journal (Refereed)
    Abstract [en]

    Benthic macrofauna is an important component linking pelagic and benthic ecosystems, especially in productive coastal areas. Through their metabolism and behaviour, benthic animals affect biogeochemical fluxes between the sediment and water column. Mechanistic models that quantify these benthic-pelagic links are imperative to understand the functioning of coastal ecosystems. In this study, we develop a dynamic model of benthic macrofauna to quantify the relationship between organic matter input and benthic macrofaunal biomass in the coastal zone. The model simulates the carbon dynamics of three functional groups of benthic macrofauna and their sediment food sources and is forced by a hydrodynamic-biogeochemical model simulating pelagic physical and biological dynamics. The model reproduces measured time-series of macrofaunal biomass from two coastal sites with contrasting sedimentation in the Baltic Sea in 1993-2005 with comparatively high accuracy, including a major increase at one of the sites dominated by the bivalve Limecola (Macoma) balthica. This shift in community composition suggests altered pathways of organic matter degradation: 39% of simulated sedimentation was mineralised by macrofauna in 2005 compared to 10% in 1995. From the early 2000s onward macrofaunal biomass seems to be food-limited, as ca 80% of organic carbon sedimentation was processed by the deposit-feeding macrofauna at both sites. This model is a first step to help quantify the role of macrofauna in marine coastal ecosystem functioning and biogeochemical cycles and build predictive capacity of the effects of anthropogenic stressors, such as eutrophication and climate change, on coastal ecosystems.

  • 6. Gammal, Johanna
    et al.
    Järnström, Marie
    Bernard, Guillaume
    Norkko, Joanna
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Environmental Context Mediates Biodiversity–Ecosystem Functioning Relationships in Coastal Soft-sediment Habitats2019In: Ecosystems (New York. Print), ISSN 1432-9840, E-ISSN 1435-0629, Vol. 22, no 1, p. 137-151Article in journal (Refereed)
    Abstract [en]

    The ongoing loss of biodiversity and global environmental changes severely affect the structure of coastal ecosystems. Consequences, in terms of ecosystem functioning, are, however, difficult to predict because the context dependency of the biodiversity–ecosystem function relationships within these heterogeneous seascapes is poorly understood. To assess the effects of biological and environmental factors in mediating ecosystem functioning (nutrient cycling) in different natural habitats, intact sediment cores were collected at 18 sites on a grain size gradient from coarse sand to silt, with varying organic matter content and vegetation. To assess ecosystem functioning, solute fluxes (O2, NH4+, PO43−, Si) across the sediment–water interface were measured. The macrofaunal communities changed along the grain size gradient with higher abundance, biomass and number of species in coarser sediments and in habitats with more vegetation. Across the whole gradient, the macrofauna cumulatively accounted for 25% of the variability in the multivariate solute fluxes, whereas environmental variables cumulatively accounted for 20%. Only the biomass and abundance of a few of the most dominant macrofauna species, not the number of species, appeared to contribute significantly to the nutrient recycling processes. Closer analyses of different sediment types (grouped into coarse, medium and fine sediment) showed that the macrofauna was an important predictor in all sediment types, but had the largest impact in fine and medium sediments. The results imply that even if the ecosystem functioning is similar in different sediment types, the underpinning mechanisms are different, which makes it challenging to generalize patterns of functioning across the heterogeneous shallow coastal zones.

  • 7.
    Griffiths, Jennifer R.
    et al.
    Stockholm University, Faculty of Science, Department of Ecology, Environment and Plant Sciences.
    Kadin, Martina
    Stockholm University, Faculty of Science, Stockholm Resilience Centre.
    Nascimento, Francisco J. A.
    Stockholm University, Faculty of Science, Department of Ecology, Environment and Plant Sciences.
    Tamelander, Tobias
    Törnroos, Anna
    Bonaglia, Stefano
    Stockholm University, Faculty of Science, Department of Geological Sciences. Lund University, Sweden.
    Bonsdorff, Erik
    Brüchert, Volker
    Stockholm University, Faculty of Science, Department of Geological Sciences.
    Gårdmark, Anna
    Järnström, Marie
    Kotta, Jonne
    Lindegren, Martin
    Nordström, Marie C.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Olsson, Jens
    Weigel, Benjamin
    Zydelis, Ramunas
    Blenckner, Thorsten
    Stockholm University, Faculty of Science, Stockholm Resilience Centre.
    Niiranen, Susa
    Stockholm University, Faculty of Science, Stockholm Resilience Centre.
    Winder, Monika
    Stockholm University, Faculty of Science, Department of Ecology, Environment and Plant Sciences.
    The importance of benthic-pelagic coupling for marine ecosystem functioning in a changing world2017In: Global Change Biology, ISSN 1354-1013, E-ISSN 1365-2486, Vol. 23, no 6, p. 2179-2196Article, review/survey (Refereed)
    Abstract [en]

    Benthic-pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic-pelagic coupling processes and their potential sensitivity to three anthropogenic pressures - climate change, nutrient loading, and fishing - using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic-pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic-pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.

  • 8. Gustafsson, Camilla
    et al.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Quantifying the importance of functional traits for primary production in aquatic plant communities2019In: Journal of Ecology, ISSN 0022-0477, E-ISSN 1365-2745, Vol. 107, no 1, p. 154-166Article in journal (Refereed)
    Abstract [en]

    1. Aquatic plant meadows are important coastal habitats that sustain many ecosystem functions such as primary production and carbon sequestration. Currently, there is a knowledge gap in understanding which plant functional traits, for example, leaf size or plant height underlie primary production in aquatic plant communities.

    2. To study how plant traits are related to primary production, we conducted a field survey in the Baltic Sea, Finland, which is characterized by high plant species and functional diversity. Thirty sites along an exposure gradient were sampled (150 plots), and nine plant morphological and chemical traits measured. The aim was to discern how community-weighted mean traits affect community production and whether this relationship changes along an environmental gradient using structural equation modelling (SEM).

    3. Plant height had a direct positive effect on production along an exposure gradient (r=0.33) and indirect effects through two leaf chemical traits, leaf delta N-15 and leaf delta C-13 (r=0.24 and 0.18, respectively) resulting in a total effect of 0.28. In plant communities experiencing varying exposure, traits such as root N concentration and leaf delta N-15 had positive and negative effects on production, respectively.

    4. Synthesis. Our results demonstrate that the relationship between aquatic plant functional traits and community production is variable and changes over environmental gradients. Plant height generally has a positive effect on community production along an exposure gradient, while the link between other traits and production changes in plant communities experiencing varying degrees of exposure. Thus, the underlying biological mechanisms influencing production differ in plant communities, emphasizing the need to resolve variability and its drivers in real-world communities. Importantly, functionally diverse plant communities sustain ecosystem functioning differently and highlight the importance of benthic diversity for coastal ecosystem stability.

  • 9.
    Humborg, Christoph
    et al.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Geibel, Marc C.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre.
    Sun, Xiaole
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre.
    McCrackin, Michelle
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre.
    Mörth, Carl-Magnus
    Stockholm University, Faculty of Science, Department of Geological Sciences.
    Stranne, Christian
    Stockholm University, Faculty of Science, Department of Geological Sciences.
    Jakobsson, Martin
    Stockholm University, Faculty of Science, Department of Geological Sciences.
    Gustafsson, Bo
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Sokolov, Alexander
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Norkko, Joanna
    High Emissions of Carbon Dioxide and Methane From the Coastal Baltic Sea at the End of a Summer Heat Wave2019In: Frontiers in Marine Science, E-ISSN 2296-7745, Vol. 6, article id 493Article in journal (Refereed)
    Abstract [en]

    The summer heat wave in 2018 led to the highest recorded water temperatures since 1926 - up to 21 degrees C - in bottom coastal waters of the Baltic Sea, with implications for the respiration patterns in these shallow coastal systems. We applied cavity ring-down spectrometer measurements to continuously monitor carbon dioxide (CO2) and methane (CH4) surface-water concentrations, covering the coastal archipelagos of Sweden and Finland and the open and deeper parts of the Northern Baltic Proper. This allowed us to (i) follow an upwelling event near the Swedish coast leading to elevated CO2 and moderate CH 4 outgassing, and (ii) to estimate CH4 sources and fluxes along the coast by investigating water column inventories and air-sea fluxes during a storm and an associated downwelling event. At the end of the heat wave, before the storm event, we found elevated CO2 (1583 mu atm) and CH4 (70 nmol/L) concentrations. During the storm, a massive CO2 sea-air flux of up to 274 mmol m(-2) d(-1) was observed. While water-column CO2 concentrations were depleted during several hours of the storm, CH4 concentrations remained elevated. Overall, we found a positive relationship between CO2 and CH4 wind-driven sea-air fluxes, however, the highest CH4 fluxes were observed at low winds whereas highest CO2 fluxes were during peak winds, suggesting different sources and processes controlling their fluxes besides wind. We applied a box-model approach to estimate the CH4 supply needed to sustain these elevated CH4 concentrations and the results suggest a large source flux of CH4 to the water column of 2.5 mmol m(-2) d(-1). These results are qualitatively supported by acoustic observations of vigorous and widespread outgassing from the sediments, with flares that could be traced throughout the water column penetrating the pycnocline and reaching the sea surface. The results suggest that the heat wave triggered CO2 and CH4 fluxes in the coastal zones that are comparable with maximum emission rates found in other hot spots, such as boreal and arctic lakes and wetlands. Further, the results suggest that heat waves are as important for CO2 and CH4 sea-air fluxes as the ice break up in spring.

  • 10. Joensuu, M.
    et al.
    Pilditch, C. A.
    Harris, R.
    Hietanen, S.
    Pettersson, H.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Sediment properties, biota, and local habitat structure explain variation in the erodibility of coastal sediments2018In: Limnology and Oceanography, ISSN 0024-3590, E-ISSN 1939-5590, Vol. 63, no 1, p. 173-186Article in journal (Refereed)
    Abstract [en]

    Sediment resuspension is a frequent phenomenon in coastal areas and a key driver for many ecosystem functions. Sediment resuspension is often linked to biological and anthropogenic activities, which in combination with hydrodynamic forcing initiate sediment erosion and resuspension, if the erosion threshold (tau(c)) is exceeded. Despite its importance to ecosystem functions very few studies have provided measurements on natural assemblages for subtidal sediments. The aim of this study was to determinate key environmental variables regulating sediment resuspension potential across a sedimentary gradient in a subtidal coastal environment. In order to explore this, we sampled 16 sites encompassing a wide variety in environmental variables (e.g., grain size distribution, macrofaunal communities, vegetation) in the Gulf of Finland, Baltic Sea. A core-based erosion device (EROMES) was used to determine sediment resuspension potential measures of erosion threshold, erosion rate (ER), and erosion constant (m(e)). Based on abiotic and biotic properties sampled, sediments diverged into two distinct groups; cohesive (muddy) and noncohesive (sandy) sediments. Results showed that abiotic sediment properties explained 38-53% and 15-36% of the total variation in resuspension potential measures in muddy and sandy sediments, respectively. In cumulative models, biota accounted for 12-26% and 6-24% to the total variation in muddy and sandy sediments, respectively. Sediment erodibility and resuspension potential of natural sediments is highly variable from local habitats to a larger seascape scale. Our results underline the importance of biota to resuspension potential measures in spatially variable environments.

  • 11. Kauppi, L.
    et al.
    Bernard, G.
    Bastrop, R.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Norkko, J.
    Increasing densities of an invasive polychaete enhance bioturbation with variable effects on solute fluxes2018In: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 8, article id 7619Article in journal (Refereed)
    Abstract [en]

    Bioturbation is a key process affecting nutrient cycling in soft sediments. The invasive polychaete genus Marenzelleria spp. has established successfully throughout the Baltic Sea increasing species and functional diversity with possible density-dependent effects on bioturbation and associated solute fluxes. We tested the effects of increasing density of M. arctia, M. viridis and M. neglecta on bioturbation and solute fluxes in a laboratory experiment. Benthic communities in intact sediment cores were manipulated by adding increasing numbers of Marenzelleria spp. The results showed that Marenzelleria spp. in general enhanced all bioturbation metrics, but the effects on solute fluxes varied depending on the solute, on the density and species identity of Marenzelleria, and on the species and functional composition of the surrounding community. M. viridis and M. neglecta were more important in predicting variation in phosphate and silicate fluxes, whereas M. arctia had a larger effect on nitrogen cycling. The complex direct and indirect pathways indicate the importance of considering the whole community and not just species in isolation in the experimental studies. Including these interactions provides a way forward regarding our understanding of the complex ecosystem effects of invasive species.

  • 12. Kauppi, L.
    et al.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Norkko, J.
    Seasonal population dynamics of the invasive polychaete genus Marenzelleria spp. in contrasting soft-sediment habitats2018In: Journal of Sea Research, ISSN 1385-1101, E-ISSN 1873-1414, Vol. 131, p. 46-60Article in journal (Refereed)
    Abstract [en]

    Three species of the invasive polychaete genus Marenzelleria are among the dominant benthic taxa in many, especially deeper, areas in the Baltic Sea. The population dynamics of the polychaetes in the Baltic are, however, still largely unknown. We conducted monthly samplings of the benthic communities and environmental parameters at five sites with differing depths and sediment characteristics in the northern Baltic Sea (59 degrees 50.896', 23 degrees 15.092') to study the population dynamics, productivity and growth of Marenzelleria spp. from April 2013 to June 2014. The species of Marenzelleria occurring at the study sites were identified by genetic analyses. At the deepest site (33 m) only M. arctia was present, while all three species were found at the shallower, muddy sites (up to 20 m depth). At the shallow (6 m) sandy site only M. viridis and M. neglecta occurred. The sites differed in the seasonal dynamics of the Marenzelleria spp. population, reflecting the different species identities. The muddy sites up to 20 m depth showed clear seasonal dynamics, with the population practically disappearing by winter, whereas more stable populations occurred at the deepest site and at the sandy site. The highest density, biomass and production were observed at the 20 m deep, organic-rich muddy site where all three species recruited. The seasonally very high densities are likely to have important consequences for organic matter processing, and species interactions at these sites. The observed high productivity of the populations has possibly facilitated their establishment, and considerably increased secondary production in especially the deeper areas.

  • 13. Kauppi, L.
    et al.
    Norkko, J.
    Ikonen, J.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Seasonal variability in ecosystem functions: quantifying the contribution of invasive species to nutrient cycling in coastal ecosystems2017In: Marine Ecology Progress Series, ISSN 0171-8630, E-ISSN 1616-1599, Vol. 572, p. 193-207Article in journal (Refereed)
    Abstract [en]

    Benthic ecosystems at temperate and high latitudes experience marked seasonal variation in the environmental factors affecting nutrient remineralization processes both directly and indirectly through their effects on the benthic communities. The invasive polychaete genus Marenzelleria represents new functionality in Baltic Sea sediments through its deep burrowing and extensive gallery formation, thus possibly greatly affecting benthic oxygen and nutrient fluxes. We assessed the seasonal contribution of Marenzelleria spp. to fluxes of solutes in monthly field measurements at 2 sites, 10 and 33 m deep, in the northern Baltic Proper over 1 yr. In general, the fluxes of inorganic nutrients and oxygen were higher during summer than during winter, and the seasonal variation was more pronounced at the deeper, more biologically active site. By using variation partitioning, we were able to demonstrate that Marenzelleria and other macrofauna could account for up to 92% of the variation in the fluxes depending on the site and season. Fauna was the most important in predicting the fluxes in spring when the sediment organic content and the abundance of juvenile Marenzelleria spp. were highest, while during e.g. winter, the influence of Marenzelleria spp., even though abundant, on solute fluxes was negligible. The results from this study have implications for management, and, importantly, for the modelling of nutrient budgets often based on values from studies conducted during the summer period only, thus possibly greatly miscalculating the annual nutrient fluxes.

  • 14. Meysick, Lukas
    et al.
    Ysebaert, Tom
    Jansson, Anna
    Montserrat, Francesc
    Valanko, Sebastian
    Villnäs, Anna
    Boström, Christoffer
    Norkko, Joanna
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Context-dependent community facilitation in seagrass meadows along a hydrodynamic stress gradient2019In: Journal of Sea Research, ISSN 1385-1101, E-ISSN 1873-1414, Vol. 150, p. 8-23Article in journal (Refereed)
    Abstract [en]

    Foundation species host diverse associated communities by ameliorating environmental stress. The strength of this facilitative effect can be highly dependent on the underlying biotic and abiotic context. We investigated community level patterns of macrofauna associated with and adjacent to the marine foundation species eelgrass (Zostera marina) along a hydrodynamic stress gradient. We could demonstrate that the relative importance of this foundation species for its infaunal community increases with environmental variables associated with increasing hydrodynamic stress (depth, sand ripples formation, sediment grain size and organic content). Faunal assemblages in proximity to the Zostera patch edges, however, showed no (infauna) or negative (epifauna) response to hydrodynamic stress. Our study highlights that the facilitative outcome of a foundation species is conditional to the faunal assemblage in question and can be highly variable even between positions within the habitat.

  • 15. Momigliano, Paolo
    et al.
    Jokinen, Henri
    Fraimout, Antoine
    Florin, Ann-Britt
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Merila, Juha
    Extraordinarily rapid speciation in a marine fish2017In: Proceedings of the National Academy of Sciences of the United States of America, ISSN 0027-8424, E-ISSN 1091-6490, Vol. 114, no 23, p. 6074-6079Article in journal (Refereed)
    Abstract [en]

    Divergent selection may initiate ecological speciation extremely rapidly. How often and at what pace ecological speciation proceeds to yield strong reproductive isolation is more uncertain. Here, we document a case of extraordinarily rapid speciation associated with ecological selection in the postglacial Baltic Sea. European flounders (Platichthys flesus) in the Baltic exhibit two contrasting reproductive behaviors: pelagic and demersal spawning. Demersal spawning enables flounders to thrive in the low salinity of the Northern Baltic, where eggs cannot achieve neutral buoyancy. We show that demersal and pelagic flounders are a species pair arising from a recent event of speciation. Despite having a parapatric distribution with extensive overlap, the two species are reciprocally monophyletic and show strongly bimodal genotypic clustering and no evidence of contemporary migration, suggesting strong reproductive isolation. Divergence across the genome is weak but shows strong signatures of selection, a pattern suggestive of a recent ecological speciation event. We propose that spawning behavior in Baltic flounders is the trait under ecologically based selection causing reproductive isolation, directly implicating a process of ecological speciation. We evaluated different possible evolutionary scenarios under the approximate Bayesian computation framework and estimate that the speciation process started in allopatry similar to 2,400 generations ago, following the colonization of the Baltic by the demersal lineage. This is faster than most known cases of ecological speciation and represents the most rapid event of speciation ever reported for any marine vertebrate.

  • 16.
    Rodil, Iván F.
    et al.
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Lucena-Moya, Paloma
    Jokinen, Henri
    Ollus, Victoria
    Wennhage, Håkan
    Villnäs, Anna
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    The role of dispersal mode and habitat specialization for metacommunity structure of shallow beach invertebrates2017In: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 12, no 2, article id e0172160Article in journal (Refereed)
    Abstract [en]

    Metacommunity ecology recognizes the interplay between local and regional patterns in contributing to spatial variation in community structure. In aquatic systems, the relative importance of such patterns depends mainly on the potential connectivity of the specific system. Thus, connectivity is expected to increase in relation to the degree of water movement, and to depend on the specific traits of the study organism. We examined the role of environmental and spatial factors in structuring benthic communities from a highly connected shallow beach network using a metacommunity approach. Both factors contributed to a varying degree to the structure of the local communities suggesting that environmental filters and dispersal-related mechanisms played key roles in determining abundance patterns. We categorized benthic taxa according to their dispersal mode (passive vs. active) and habitat specialization (generalist vs. specialist) to understand the relative importance of environment and dispersal related processes for shallow beach metacommunities. Passive dispersers were predicted by a combination of environmental and spatial factors, whereas active dispersers were not spatially structured and responded only to local environmental factors. Generalists were predicted primarily by spatial factors, while specialists were only predicted by local environmental factors. The results suggest that the role of the spatial component in metacommunity organization is greater in open coastal waters, such as shallow beaches, compared to less-connected environmentally controlled aquatic systems. Our results also reveal a strong environmental role in structuring the benthic metacommunity of shallow beaches. Specifically, we highlight the sensitivity of shallow beach macrofauna to environmental factors related to eutrophication proxies.

  • 17. Snelgrove, Paul V. R.
    et al.
    Soetaert, Karline
    Solan, Martin
    Thrush, Simon
    Wei, Chih-Lin
    Danovaro, Roberto
    Fulweiler, Robinson W.
    Kitazato, Hiroshi
    Ingole, Baban
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Parkes, R. John
    Volkenborn, Nils
    Global Carbon Cycling on a Heterogeneous Seafloor2018In: Trends in Ecology & Evolution, ISSN 0169-5347, E-ISSN 1872-8383, Vol. 33, no 2, p. 96-105Article, review/survey (Refereed)
    Abstract [en]

    Diverse biological communities mediate the transformation, transport, and storage of elements fundamental to life on Earth, including carbon, nitrogen, and oxygen. However, global biogeochemical model outcomes can vary by orders of magnitude, compromising capacity to project realistic ecosystem responses to planetary changes, including ocean productivity and climate. Here, we compare global carbon turnover rates estimated using models grounded in biological versus geochemical theory and argue that the turnover estimates based on each perspective yield divergent outcomes. Importantly, empirical studies that include sedimentary biological activity vary less than those that ignore it. Improving the relevance of model projections and reducing uncertainty associated with the anticipated consequences of global change requires reconciliation of these perspectives, enabling better societal decisions on mitigation and adaptation.

  • 18. Villnas, Anna
    et al.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Lehtonen, Kari K.
    Multi-level responses of Macoma balthica to recurring hypoxic disturbance2019In: Journal of Experimental Marine Biology and Ecology, ISSN 0022-0981, E-ISSN 1879-1697, Vol. 510, p. 64-72Article in journal (Refereed)
    Abstract [en]

    The frequency of seasonal and short-term hypoxia is increasing in coastal seas. How such repeated disturbances affect key species that have important roles for ecosystem processes and functions remains, however, unknown. By performing a field experiment we explored if the bivalve Macoma balthica can cope with short-term, recurring hypoxic stress, and investigated how hypoxia affects the condition of surviving bivalves. By combining data on different levels of biological organization, i.e., on physiology (biomarker response), behaviour and demography, we identified stress responses before the population declined. One pulse of hypoxic disturbance (3 days) resulted in behavioural alterations, as adult M. balthica extended their siphons, emerged towards the sediment surface and expressed decreased reburial rates. However, the demographic structure of the population remained unaltered. Several pulses of recurring hypoxic stress resulted in physiological response with changes in glutathione reductase and acetylcholinesterase enzyme activities. The recurring hypoxic disturbance was observed to affect juvenile bivalves before adults, while pro-longed hypoxia reduced the entire bivalve population. Our results clearly show that hypoxic stress changes the behaviour and physiology of M. balthica before demographic changes occur, which is likely to have severe implications for the contribution of this key species to ecosystem functioning. That a combination of measures at different levels of organization can detect disturbances at an early stage suggests that such an approach would be useful for assessing the effects of disturbances on marine ecosystems that are increasingly affected by anthropogenic change.

  • 19. Villnäs, Anna
    et al.
    Hewitt, Judi
    Snickars, Martin
    Westerbom, Mats
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Template for using biological trait groupings when exploringlarge-scale variation in seafloor multifunctionality2018In: Ecological Applications, ISSN 1051-0761, E-ISSN 1939-5582, Vol. 28, no 1, p. 78-94Article in journal (Refereed)
    Abstract [en]

    Understanding large-scale spatial variation in ecosystem properties and associated functionality is key for successful conservation of ecosystems. This study provides a template for how to estimate differences in ecosystem functionality over large spatial scales by using groupings of biological traits. We focus on trait groupings that describe three important benthic ecosystem properties, namely bioturbation, community stability, and juvenile dispersal. Recognizing that groups of traits interact and are constrained within an organism, we statistically define important functional trait subgroups that describe each ecosystem property. The sub-groups are scored according to their weighted ecological impact to gain an overall estimation of the cumulative expression of each ecosystem property at individual sites. Furthermore, by assigning each property a value relative to its observed maximum, and by summing up the individual property values, we offer an estimate of benthic ecosystem multi-functionality. Based on a spatially extensive benthic data set, we were able to identify coastal areas with high and low potential for the considered benthic ecosystem properties and the measure of ecosystem multifunctionality. Importantly, we show that a large part of the spatial variation in functional trait sub-groups and in benthic ecosystem multifunctionality was explained by environmental change. Our results indicate that through this simplification it is possible to estimate the functionality of the seafloor. Such information is vital in marine spatial planning efforts striving to balance the utilization with the preservation of natural resources.

  • 20. Virta, Leena
    et al.
    Gammal, Hanna
    Järnström, Marie
    Bernard, Guillaume
    Soininen, Janne
    Norkko, Joanna
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    The diversity of benthic diatoms affects ecosystem productivity in heterogeneous coastal environments2019In: Ecology, ISSN 0012-9658, E-ISSN 1939-9170Article in journal (Refereed)
    Abstract [en]

    The current decrease in biodiversity affects all ecosystems, and the impacts of diversity on ecosystem functioning need to be resolved. So far, marine studies about diversity-ecosystem productivity-relationships have concentrated on small-scale, controlled experiments, with often limited relevance to natural ecosystems. Here, we provide a real-world study on the effects of microorganismal diversity (measured as the diversity of benthic diatom communities) on ecosystem productivity (using chlorophyll a concentration as a surrogate) in a heterogeneous marine coastal archipelago. We collected 78 sediment cores at 17 sites in the northern Baltic Sea and found exceptionally high diatom diversity (328 observed species). We used structural equation models and quantile regression to explore relationships between diatom diversity and productivity. Previous studies have found contradictory results in the relationship between microorganismal diversity and ecosystem productivity, but we showed a linear and positive basal relationship between diatom diversity and productivity, which indicates that diatom diversity most likely forms the lowest boundary for productivity. Thus, although productivity can be high even when diatom diversity is low, high diatom diversity supports high productivity. The trait composition was more effective than taxonomical composition in showing such a relationship, which could be due to niche complementarity. Our results also indicated that environmental heterogeneity leads to substantial patchiness in the diversity of benthic diatom communities, mainly induced by the variation in sediment organic matter content. Therefore, future changes in precipitation and river runoff and associated changes in the quality and quantity of organic matter in the sea, will also affect diatom communities and, hence, ecosystem productivity. Our study suggests that benthic microorganisms are vital for ecosystem productivity, and together with the substantial heterogeneity of coastal ecosystems, they should be considered when evaluating the potential productivity of coastal areas.

  • 21. Virtanen, Elina A.
    et al.
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. University of Helsinki, Finland.
    Nyström Sandman, Antonia
    Viitasalo, Markku
    Identifying areas prone to coastal hypoxia - the role of topography2019In: Biogeosciences, ISSN 1726-4170, E-ISSN 1726-4189, Vol. 16, no 16, p. 3183-3195Article in journal (Refereed)
    Abstract [en]

    Hypoxia is an increasing problem in marine ecosystems around the world. While major advances have been made in our understanding of the drivers of hypoxia, challenges remain in describing oxygen dynamics in coastal regions. The complexity of many coastal areas and lack of detailed in situ data have hindered the development of models describing oxygen dynamics at a sufficient spatial resolution for efficient management actions to take place. It is well known that the enclosed nature of seafloors and reduced water mixing facilitates hypoxia formation, but the degree to which topography contributes to hypoxia formation and small-scale variability of coastal hypoxia has not been previously quantified. We developed simple proxies of seafloor heterogeneity and modeled oxygen deficiency in complex coastal areas in the northern Baltic Sea. According to our models, topographical parameters alone explained similar to 80 % of hypoxia occurrences. The models also revealed that less than 25 % of the studied seascapes were prone to hypoxia during late summer (August-September). However, large variation existed in the spatial and temporal patterns of hypoxia, as certain areas were prone to occasional severe hypoxia (O-2 < 2 mg L-1), while others were more susceptible to recurrent moderate hypoxia (O-2 < 4.6 mg L-1). Areas identified as problematic in our study were characterized by low exposure to wave forcing, high topographic shelter from surrounding areas and isolation from the open sea, all contributing to longer water residence times in seabed depressions. Deviations from this topographical background are probably caused by strong currents or high nutrient loading, thus improving or worsening oxygen status, respectively. In some areas, connectivity with adjacent deeper basins may also influence coastal oxygen dynamics. Developed models could boost the performance of biogeochemical models, aid developing nutrient abatement measures and pinpoint areas where management actions are most urgently needed.

  • 22. Westerbom, Mats
    et al.
    Lappalainen, Antti
    Mustonen, Olli
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. Tvärminne Zoological Station, Finland.
    Trophic overlap between expanding and contracting fish predators in a range margin undergoing change2018In: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 8, article id 7895Article in journal (Refereed)
    Abstract [en]

    Climate change is predicted to cause a freshening of the Baltic Sea, facilitating range expansions of freshwater species and contractions of marine. Resident marine flounders (Platichthys flesus) and expansive freshwater roach (Rutilus rutilus) are dominant consumers in the Baltic Sea sublittoral where they occur in partial sympatry. By comparing patterns of resource use by flounders and roach along a declining resource gradient of blue mussels (Mytilus trossulus) our aim was to explore predator functional responses and the degree of trophic overlap. Understanding the nature of density-dependent prey acquisition has important implications for predicting population dynamics of both predators and their shared prey. Results showed a highly specialized diet for both species, high reliance on blue mussels throughout the range, similar prey size preference and high trophic overlap. Highest overlap occurred where blue mussels were abundant but overlap was also high where they were scarce. Our results highlight the importance of a single food item - the blue mussel - for both species, likely promoting high population size and range expansion of roach. Findings also suggest that range expansion of roach may have a top-down structuring force on mussels that differ in severity and location from that originating from resident flounders.

  • 23. Westerbom, Mats
    et al.
    Mustonen, Olli
    Jaatinen, Kim
    Kilpi, Mikael
    Norkko, Alf
    Stockholm University, Faculty of Science, Stockholm University Baltic Sea Centre. Helsinki University, Finland.
    Population Dynamics at the Range Margin: Implications of Climate Change on Sublittoral Blue Mussels (Mytilus trossulus)2019In: Frontiers in Marine Science, E-ISSN 2296-7745, Vol. 6, article id UNSP 292Article in journal (Refereed)
    Abstract [en]

    Examining changes in abundance and demographic rates at species distribution margins may provide the first signs of broader species responses to environmental change. Still, the joint impact of space and time have remained relatively unstudied in most marginal regions. In order to examine the influence of climate variability on mussel distribution patterns, we monitored three sublittoral and marginal blue mussel (Mytilus trossulus) populations, spaced along a salinity gradient. Densities and biomasses peaked toward the saltier parts of the study area and showed relatively larger variations toward the low saline edge. Temporally, the areas showed a consistent increase in abundance after a synchronized large-scale recruitment event, which was followed by a decline in population size, occurring much faster toward the very range edge. Salinity, temperature, winter severity, and wave exposure explained most of the spatiotemporal variation in mussel abundances and adults showed positive effects on recruit abundance. We show empirically that the dynamics of edge populations are not driven by large changes in climate variables but that small spatial and temporal changes in key environmental variables have large and non-linear population level effects. Our results also show that fluctuating recruitment is a key factor for population stability affecting the storage potential of marginal populations, which dramatically decrease toward the edge. Our study provides a window into future population patterns and processes that drive marginal mussel populations in an altered sea characterized by rising temperature and declining salinity.

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