Dataset from a mesocosm experiment on brownification in the Baltic Sea.

Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment over 15 days to test the effect of this on the planktonic ecosystem. The experiment was set up in 2.2 m3 plastic bags moored outside the Tvärminne Zoological Station at the SW coast of Finland. We used four treatments, each with three replicates: control (Contr) without any manipulation; addition of a commercially available organic carbon additive called HuminFeed (Hum; 2 mg L-1); addition of inorganic nutrients (Nutr; 5.7 µM NH4 and 0.65µM PO4); and a final treatment of combined Nutr and Hum (Nutr+Hum) additions. Water samples were taken daily, and measured variables included water transparency, organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry.

Brownification affects phytoplankton community composition but not primary productivity in eutrophic coastal waters: A mesocosm experiment in the Baltic Sea.

Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment (15 d) to test the effect of this on the planktonic ecosystem expecting reduced primary production and shifts in the phytoplankton community composition. The experiment was set up in 2.2 m3 mesocosm bags using four treatments, each with three replicates: control (Contr) without any manipulation, organic carbon additive HuminFeed (Hum; 2 mg L-1), inorganic nutrients (Nutr; 5.7 µM NH4 and 0.65 µM PO4), and combined Nutr and Hum (Nutr + Hum) additions. Measured variables included organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry. The bags with added inorganic nutrients developed a phytoplankton bloom that depleted inorganic N at day 6, followed by a rapid decline in Chla. Brownification did not reduce primary production at the tested concentration. Bacterial production was lowest in the Contr, but similar in the three treatments receiving additions likely due to increased carbon available for heterotrophic bacteria. Picoeukaryotes clearly benefited by brownification after inorganic N depletion, which could be due to more effective nutrient recycling, nutrient affinity, light absorption, or alternatively lower grazing pressure. In conclusion, brownification shifted the phytoplankton community composition towards smaller species with potential effects on carbon fluxes, such as sinking rates and export to the sea floor.

Effects of Added Humic Substances and Nutrients on Photochemical Degradation of Dissolved Organic Matter in a Mesocosm Amendment Experiment in the Gulf of Finland, Baltic Sea.

Humic substances, a component of terrestrial dissolved organic matter (tDOM), contribute to dissolved organic matter (DOM) and chromophoric DOM (CDOM) in coastal waters, and have significant impacts on biogeochemistry. There are concerns in recent years over browning effects in surface waters due to increasing tDOM inputs, and their negative impacts on aquatic ecosystems, but relatively little work has been published on estuaries and coastal waters. Photodegradation could be a significant sink for tDOM in coastal environments, but the rates and efficiencies are poorly constrained. We conducted large-scale DOM photodegradation experiments in mesocosms amended with humic substances and nutrients in the Gulf of Finland to investigate the potential of photochemistry to remove added tDOM and the interactions of DOM photochemistry with eutrophication. The added tDOM was photodegraded rapidly, as CDOM absorption decreased and spectral slopes increased with increasing photons absorbed in laboratory experiments. The in situ DOM optical properties became similar among the control, humic- and humic+nutrients-amended mesocosm samples toward the end of the amendment experiment, indicating degradation of the excess CDOM/DOM through processes including photodegradation. Nutrient additions did not significantly influence the effects of added humic substances on CDOM optical property changes, but induced changes in DOM removal.

Does the use of biological traits predict a smooth landscape of ecosystem functioning?

The biodiversity crisis has increased interest in understanding the role of biodiversity for ecosystem functioning. Functional traits are often used to infer ecosystem functions to increase our understanding of these relationships over larger spatial scales. The links between specific traits and ecosystem functioning are, however, not always well established. We investigated how the choice of analyzing either individual species, selected modalities, or trait combinations affected the spatial patterns observed on a sandflat and how this was related to the natural variability in ecosystem functioning. A large dataset of 400 benthic macrofauna samples was used to explore distribution patterns. We hypothesized that (1) if multiple species (redundancy) represent a trait combination or a modality their spatial patterns would be smoothed out, and (2) the lost spatial variability within a trait combination or modality, due to the smoothing effect, would potentially affect their utility for predicting ecosystem functioning (tested on a dataset of 24 samples). We predicted that species would show heterogeneous small spatial patterns, while modalities and trait combinations would show larger and more homogeneous patterns because they would represent a collection of many distributions. If modalities and trait combinations are better predictors of ecosystem functioning than species, then the smoother spatial patterns of modalities and trait combinations would result in a more homogeneous landscape of ecosystem function and the number of species exhibiting specific traits would provide functional redundancy. Our results showed some smoothing of spatial patterns progressing from species through modalities to trait combinations, but generally spatial patterns reflected a few dominant key species. Moreover, some individual modalities and species explained more or equal proportions of the variance in the ecosystem functioning than the combined traits. The findings thus suggest that only some spatial variability is lost when species are combined into modalities and trait combinations and that a homogeneous landscape of ecosystem function is not likely.

Monitoring of ticks and tick-borne pathogens through a nationwide research station network in Finland.

In 2015 a long-term, nationwide tick and tick-borne pathogen (TBP) monitoring project was started by the Finnish Tick Project and the Finnish Research Station network (RESTAT), with the goal of producing temporally and geographically extensive data regarding exophilic ticks in Finland. In the current study, we present results from the first four years of this collaboration. Ticks were collected by cloth dragging from 11 research stations across Finland in May-September 2015-2018 (2012-2018 in Seili). Collected ticks were screened for twelve different pathogens by qPCR: Borrelia afzelii, Borrelia garinii, Borrelia valaisiana, Borrelia burgdorferi sensu stricto, Borrelia miyamotoi, Babesia spp., Anaplasma phagocytophilum, Rickettsia spp., Candidatus Neoehrlichia mikurensis, Francisella tularensis, Bartonella spp. and tick-borne encephalitis virus (TBEV). Altogether 15 067 Ixodes ricinus and 46 Ixodes persulcatus were collected during 68 km of dragging. Field collections revealed different seasonal activity patterns for the two species. The activity of I. persulcatus adults (only one nymph detected) was unimodal, with activity only in May-July, whereas Ixodes ricinus was active from May to September, with activity peaks in September (nymphs) or July-August (adults). Overall, tick densities were higher during the latter years of the study. Borrelia burgdorferi sensu lato were the most common pathogens detected, with 48.9 ±â€¯8.4% (95% Cl) of adults and 25.3 ±â€¯4.4% of nymphs carrying the bacteria. No samples positive for F. tularensis, Bartonella or TBEV were detected. This collaboration project involving the extensive Finnish Research Station network has ensured enduring and spatially extensive, long-term tick data collection to the foreseeable future.

Increasing oxygen deficiency changes rare and moderately abundant bacterial communities in coastal soft sediments.

Coastal hypoxia is a major environmental problem worldwide. Hypoxia-induced changes in sediment bacterial communities harm marine ecosystems and alter biogeochemical cycles. Nevertheless, the resistance of sediment bacterial communities to hypoxic stress is unknown. We investigated changes in bacterial communities during hypoxic-anoxic disturbance by artificially inducing oxygen deficiency to the seafloor for 0, 3, 7, and 48 days, with subsequent molecular biological analyses. We further investigated relationships between bacterial communities, benthic macrofauna and nutrient effluxes across the sediment-water-interface during hypoxic-anoxic stress, considering differentially abundant operational taxonomic units (OTUs). The composition of the moderately abundant OTUs changed significantly after seven days of oxygen deficiency, while the abundant and rare OTUs first changed after 48 days. High bacterial diversity maintained the resistance of the communities during oxygen deficiency until it dropped after 48 days, likely due to anoxia-induced loss of macrofaunal diversity and bioturbation. Nutrient fluxes, especially ammonium, correlated positively with the moderate and rare OTUs, including potential sulfate reducers. Correlations may reflect bacteria-mediated nutrient effluxes that accelerate eutrophication. The study suggests that even slightly higher bottom-water oxygen concentrations, which could sustain macrofaunal bioturbation, enable bacterial communities to resist large compositional changes and decrease the harmful consequences of hypoxia in marine ecosystems.

The diversity of benthic diatoms affects ecosystem productivity in heterogeneous coastal environments.

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.

Towards a sampling design for characterizing habitat-specific benthic biodiversity related to oxygen flux dynamics using Aquatic Eddy Covariance.

The Aquatic Eddy Covariance (AEC) technique has emerged as an important method to quantify in situ seafloor metabolism over large areas of heterogeneous benthic communities, enabling cross-habitat comparisons of seafloor productivity. However, the lack of a corresponding sampling protocol to perform biodiversity comparisons across habitats is impeding a full assessment of marine ecosystem metabolism. Here, we study a range of coastal benthic habitats, from rocky-bed communities defined by either perennial macroalgae or blue mussel beds to soft-sediment communities comprised of either seagrass, patches of different macrophyte species or bare sand. We estimated that the maximum contribution to the AEC metabolic flux can be found for a seafloor area of approximately 80 m2 with a 5 meter upstream distance of the instrument across all the habitats. We conducted a sampling approach to characterize and quantify the dominant features of biodiversity (i.e., community biomass) within the main seafloor area of maximum metabolic contribution (i.e., gross primary production and community respiration) measured by the AEC. We documented a high biomass contribution of the macroalgal Fucus vesiculosus, the seagrass Zostera marina and the macroinvertebrate Mytilus edulis to the net ecosystem metabolism of the habitats. We also documented a significant role of the bare sediments for primary productivity compared to vegetated canopies of the soft sediments. The AEC also provided insight into dynamic short-term drivers of productivity such as PAR availability and water flow velocity for the productivity estimate. We regard this study as an important step forward, setting a framework for upcoming research focusing on linking biodiversity metrics and AEC flux measurements across habitats.

Feeding type affects microplastic ingestion in a coastal invertebrate community.

Marine litter is one of the problems marine ecosystems face at present, coastal habitats and food webs being the most vulnerable as they are closest to the sources of litter. A range of animals (bivalves, free swimming crustaceans and benthic, deposit-feeding animals), of a coastal community of the northern Baltic Sea were exposed to relatively low concentrations of 10 µm microbeads. The experiment was carried out as a small scale mesocosm study to mimic natural habitat. The beads were ingested by all animals in all experimental concentrations (5, 50 and 250 beads mL(-1)). Bivalves (Mytilus trossulus, Macoma balthica) contained significantly higher amounts of beads compared with the other groups. Free-swimming crustaceans ingested more beads compared with the benthic animals that were feeding only on the sediment surface. Ingestion of the beads was concluded to be the result of particle concentration, feeding mode and the encounter rate in a patchy environment.

Post-settlement dispersal: the neglected link in maintenance of soft-sediment biodiversity.

Seafloor integrity is threatened by disturbances owing to human activities. The capacity of the system to recover from disturbances, as well as maintain resilience and function, depends on dispersal. In soft-sediment systems, dispersal continues after larval settlement, but there are very few measurements of how far the post-settlers disperse in nature. Spatial scales of post-settlement dispersal are, however, likely to be similar to pelagic larval dispersal because of continued, frequent, small-scale dispersal over longer periods. The consequences of this dispersal may be more important for the maintenance of biodiversity and metacommunity dynamics than is pelagic larval dispersal, because of the greater size and competency of the dispersers. We argue that an increased empirical understanding of post-settlement dispersal processes is key for predicting how benthic communities will respond to local disturbances and shrinking regional species pools, with implications for monitoring, managing and conserving biodiversity.

Hypoxia in the Baltic Sea: biogeochemical cycles, benthic fauna, and management.

Hypoxia has occurred intermittently over the Holocene in the Baltic Sea, but the recent expansion from less than 10 000 km(2) before 1950 to >60 000 km(2) since 2000 is mainly caused by enhanced nutrient inputs from land and atmosphere. With worsening hypoxia, the role of sediments changes from nitrogen removal to nitrogen release as ammonium. At present, denitrification in the water column and sediments is equally important. Phosphorus is currently buried in sediments mainly in organic form, with an additional contribution of reduced Fe-phosphate minerals in the deep anoxic basins. Upon the transition to oxic conditions, a significant proportion of the organic phosphorus will be remineralized, with the phosphorus then being bound to iron oxides. This iron-oxide bound phosphorus is readily released to the water column upon the onset of hypoxia again. Important ecosystems services carried out by the benthic fauna, including biogeochemical feedback-loops and biomass production, are also lost with hypoxia. The results provide quantitative knowledge of nutrient release and recycling processes under various environmental conditions in support of decision support tools underlying the Baltic Sea Action Plan.

The role of recurrent disturbances for ecosystem multifunctionality.

Ecosystem functioning is threatened by an increasing number of anthropogenic stressors, creating a legacy of disturbance that undermines ecosystem resilience. However, few empirical studies have assessed to what extent an ecosystem can tolerate repeated disturbances and sustain its multiple functions. By inducing increasingly recurring hypoxic disturbances to a sedimentary ecosystem, we show that the majority of individual ecosystem functions experience gradual degradation patterns in response to repetitive pulse disturbances. The degradation in overall ecosystem functioning was, however, evident at an earlier stage than for single ecosystem functions and was induced after a short pulse of hypoxia (i.e., three days), which likely reduced ecosystem resistance to further hypoxic perturbations. The increasing number of repeated pulse disturbances gradually moved the system closer to a press response. In addition to the disturbance regime, the changes in benthic trait composition as well as habitat heterogeneity were important for explaining the variability in overall ecosystem functioning. Our results suggest that disturbance-induced responses across multiple ecosystem functions can serve as a warning signal for losses of the adaptive capacity of an ecosystem, and might at an early stage provide information to managers and policy makers when remediation efforts should be initiated.

Size matters: implications of the loss of large individuals for ecosystem function.

Size is a fundamental organismal trait and an important driver of ecosystem functions. Although large individuals may dominate some functions and provide important habitat structuring effects, intra-specific body size effects are rarely investigated in the context of BEF relationships. We used an in situ density manipulation experiment to explore the contribution of large, deep-burrowing bivalves to oxygen and nutrient fluxes across the sediment-water interface. By manipulating bivalve size structure through the removal of large individuals, we held species identity constant, but altered the trait characteristics of the community. The number of large bivalves was the best predictor of ecosystem functioning. Our results highlight that (a) accounting for body size provides important insights into the mechanisms underpinning biodiversity effects on ecosystem function, and (b) if local disturbances are recurrent, preventing individuals from reaching large sizes, the contribution of large adults may be lost, with largely unknown implications for ecosystem functionality.

Effects of Reduced pH on Macoma balthica Larvae from a System with Naturally Fluctuating pH-Dynamics.

Ocean acidification is causing severe changes in the inorganic carbon balance of the oceans. The pH conditions predicted for the future oceans are, however, already regularly occurring in the Baltic Sea, and the system might thus work as an analogue for future ocean acidification scenarios. The characteristics of the Baltic Sea with low buffering capacity and large natural pH fluctuations, in combination with multiple other stressors, suggest that OA effects may be severe, but remain largely unexplored. A calcifying species potentially affected by low pH conditions is the bivalve Macoma balthica (L.). We investigated larval survival and development of M. balthica by exposing the larvae to a range of pH levels: 7.2, 7.4, 7.7 and 8.1 during 20 days in order to learn what the effects of reduced pH are on the larval biology and thus also potentially for the population dynamics of this key species. We found that even a slight pH decrease causes significant negative changes during the larval phase, both by slowing growth and by decreasing survival. The growth was slower in all reduced pH treatments compared to the control treatment. The size of 250 µm that is considered indicative to imminent settling in our system was reached by 22% of the larvae grown in control conditions after 20 days, whereas in all reduced pH treatments the size of 250 µm was reached by only 7-14%. The strong impact of ocean acidification on larvae is alarming as slowly growing individuals are exposed to higher predation risk in response to the longer time they are required to spend in the plankton, further decreasing the ecological competence of the species.

Consequences of increasing hypoxic disturbance on benthic communities and ecosystem functioning.

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH(4)(+) and dissolved Si. Although effluxes of PO(4)(3-) were not altered significantly, changes were observed in sediment PO(4)(3-) sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.

Ocean acidification at high latitudes: potential effects on functioning of the Antarctic bivalve Laternula elliptica.

Ocean acidification is a well recognised threat to marine ecosystems. High latitude regions are predicted to be particularly affected due to cold waters and naturally low carbonate saturation levels. This is of concern for organisms utilising calcium carbonate (CaCO(3)) to generate shells or skeletons. Studies of potential effects of future levels of pCO(2) on high latitude calcifiers are at present limited, and there is little understanding of their potential to acclimate to these changes. We describe a laboratory experiment to compare physiological and metabolic responses of a key benthic bivalve, Laternula elliptica, at pCO(2) levels of their natural environment (430 µatm, pH 7.99; based on field measurements) with those predicted for 2100 (735 µatm, pH 7.78) and glacial levels (187 µatm, pH 8.32). Adult L. elliptica basal metabolism (oxygen consumption rates) and heat shock protein HSP70 gene expression levels increased in response both to lowering and elevation of pH. Expression of chitin synthase (CHS), a key enzyme involved in synthesis of bivalve shells, was significantly up-regulated in individuals at pH 7.78, indicating L. elliptica were working harder to calcify in seawater undersaturated in aragonite (Ω(Ar) = 0.71), the CaCO(3) polymorph of which their shells are comprised. The different response variables were influenced by pH in differing ways, highlighting the importance of assessing a variety of factors to determine the likely impact of pH change. In combination, the results indicate a negative effect of ocean acidification on whole-organism functioning of L. elliptica over relatively short terms (weeks-months) that may be energetically difficult to maintain over longer time periods. Importantly, however, the observed changes in L. elliptica CHS gene expression provides evidence for biological control over the shell formation process, which may enable some degree of adaptation or acclimation to future ocean acidification scenarios.

Forecasting the limits of resilience: integrating empirical research with theory.

Despite the increasing evidence of drastic and profound changes in many ecosystems, often referred to as regime shifts, we have little ability to understand the processes that provide insurance against such change (resilience). Modelling studies have suggested that increased variance may foreshadow a regime shift, but this requires long-term data and knowledge of the functional links between key processes. Field-based research and ground-truthing is an essential part of the heuristic that marries theoretical and empirical research, but experimental studies of resilience are lagging behind theory, management and policy requirements. Empirically, ecological resilience must be understood in terms of community dynamics and the potential for small shifts in environmental forcing to break the feedbacks that support resilience. Here, we integrate recent theory and empirical data to identify ways we might define and understand potential thresholds in the resilience of nature, and thus the potential for regime shifts, by focusing on the roles of strong and weak interactions, linkages in meta-communities, and positive feedbacks between these and environmental drivers. The challenge to theoretical and field ecologists is to make the shift from hindsight to a more predictive science that is able to assist in the implementation of ecosystem-based management.