How oxygen deficiency in the Baltic Sea proper has spread and worsened: The role of ammonium and hydrogen sulphide.

Even large inflows of oxygen-rich seawater to the Baltic Proper have in recent decades given only short-lived relief from oxygen deficiency below the halocline. We analyse long-term changes in oxygen deficiency, and calculate the "total oxygen debt" [Formula: see text]OD, the oxygen required to oxidize the hydrogen sulphide (H2S) and ammonium (NH4) that builds up during stagnation periods. Since the early 1990s, oxygen below 65m has gradually decreased during successive stagnation periods, and the [Formula: see text]OD has increased, with NH4 more important than previously recognised. After the major inflow in 2014, the Baltic Proper [Formula: see text]OD has reached its highest level so far. The gradual shift of the [Formula: see text]OD to shallower sub-halocline waters in the western and northern basins has increased the risk of periodic coastal hypoxia and export of hypoxic water to the Bothnian Sea. The potential for inflows large enough to more than eliminate the [Formula: see text]OD seems limited in the near term.

Cyanobacterial blooms in the Baltic Sea: Correlations with environmental factors.

Massive cyanobacteria blooms occur almost every summer in the Baltic Sea but the capability to quantitatively predict their extent and intensity is poorly developed. Here we analyse statistical relationships between multi-decadal satellite-derived time series of the frequency of cyanobacteria surface accumulations (FCA) in the central Baltic Sea Proper and a suite of environmental variables. Over the decadal scale (∼5-20 years) FCA was highly correlated (R2 ∼ 0.69) with a set of biogeochemical variables related to the amount of phosphorus and hypoxia in bottom layers. Water temperature in the surface layer was also positively correlated with FCA at the decadal scale. In contrast, the inter-annual variations in FCA had no correlation with the biogeochemical variables. Instead, significant correlations were found with the solar shortwave direct flux in July and the sea-surface temperature, also in July. It thus appears that it is not possible to predict inter-annual fluctuations in cyanobacteria blooms from water chemistry. Moreover, environmental variables could only explain about 45% of the inter-annual variability in FCA, probably because year-to-year variations in FCA are significantly influenced by biological interactions.

Unexplained interannual oscillations of cyanobacterial blooms in the Baltic Sea.

Population oscillations in multi-species or even single species systems are well-known but have rarely been detected at the lower trophic levels in marine systems. Nitrogen fixing cyanobacteria are a major component of the Baltic Sea ecosystem and sometimes form huge surface accumulations covering most of the sea surface. By analysing a satellite-derived 39-year (1979-2017) data archive of surface cyanobacteria concentrations we have found evidence of strikingly regular interannual oscillations in cyanobacteria concentrations in the northern Baltic Sea. These oscillations have a period of ~3 years with a high-concentration year generally followed by one or two low-concentration years. Changes in abiotic factors known to influence the growth and survival of cyanobacteria could not provide an explanation for the oscillations. We therefore assume that these oscillations are intrinsic to the marine system, caused by an unknown, probably mainly biological mechanism that may be triggered by a combination of environmental factors. Interactions between different life cycle stages of cyanobacteria as well as between predator-prey or host-parasite are possible candidates for causing the oscillations.

Human activities and climate variability drive fast-paced change across the world's estuarine-coastal ecosystems.

Time series of environmental measurements are essential for detecting, measuring and understanding changes in the Earth system and its biological communities. Observational series have accumulated over the past 2-5 decades from measurements across the world's estuaries, bays, lagoons, inland seas and shelf waters influenced by runoff. We synthesize information contained in these time series to develop a global view of changes occurring in marine systems influenced by connectivity to land. Our review is organized around four themes: (i) human activities as drivers of change; (ii) variability of the climate system as a driver of change; (iii) successes, disappointments and challenges of managing change at the sea-land interface; and (iv) discoveries made from observations over time. Multidecadal time series reveal that many of the world's estuarine-coastal ecosystems are in a continuing state of change, and the pace of change is faster than we could have imagined a decade ago. Some have been transformed into novel ecosystems with habitats, biogeochemistry and biological communities outside the natural range of variability. Change takes many forms including linear and nonlinear trends, abrupt state changes and oscillations. The challenge of managing change is daunting in the coastal zone where diverse human pressures are concentrated and intersect with different responses to climate variability over land and over ocean basins. The pace of change in estuarine-coastal ecosystems will likely accelerate as the human population and economies continue to grow and as global climate change accelerates. Wise stewardship of the resources upon which we depend is critically dependent upon a continuing flow of information from observations to measure, understand and anticipate future changes along the world's coastlines.

Do deposit-feeders compete? Isotopic niche analysis of an invasion in a species-poor system.

Successful establishment of invasive species is often related to the existence of vacant niches. Competition occurs when invaders use the same limiting resources as members of the recipient community, which will be reflected in some overlap of their trophic niches. The concept of isotopic niche has been used to study trophic niche partitioning among species. Here, we present a two-year field study comparing isotopic niches of the deposit-feeding community in a naturally species-poor system. The isotopic niche analyses showed no overlap between a recent polychaete invader and any of the native species suggesting that it has occupied a vacant niche. Its narrow isotopic niche suggests specialized feeding, however, the high δ(15)N values compared to natives are most likely due to isotope fractionation effects related to nitrogen recycling and a mismatch between biological stoichiometry of the polychaete and the sediment nitrogen content. Notably, highly overlapping isotopic niches were inferred for the native species, which is surprising in a food-limited system. Therefore, our results demonstrate that invaders may broaden the community trophic diversity and enhance resource utilization, but also raise questions about the congruence between trophic and isotopic niche concepts and call for careful examination of assumptions underlying isotopic niche interpretation.

Projected future climate change and Baltic Sea ecosystem management.

Climate change is likely to have large effects on the Baltic Sea ecosystem. Simulations indicate 2-4 °C warming and 50-80 % decrease in ice cover by 2100. Precipitation may increase ~30 % in the north, causing increased land runoff of allochthonous organic matter (AOM) and organic pollutants and decreased salinity. Coupled physical-biogeochemical models indicate that, in the south, bottom-water anoxia may spread, reducing cod recruitment and increasing sediment phosphorus release, thus promoting cyanobacterial blooms. In the north, heterotrophic bacteria will be favored by AOM, while phytoplankton production may be reduced. Extra trophic levels in the food web may increase energy losses and consequently reduce fish production. Future management of the Baltic Sea must consider the effects of climate change on the ecosystem dynamics and functions, as well as the effects of anthropogenic nutrient and pollutant load. Monitoring should have a holistic approach, encompassing both autotrophic (phytoplankton) and heterotrophic (e.g., bacterial) processes.

Baltic Sea management: Successes and failures.

Severe environmental problems documented in the Baltic Sea in the 1960s led to the 1974 creation of the Helsinki Convention for the Protection of the Marine Environment of the Baltic Sea Area. We introduce this special issue by briefly summarizing successes and failures of Baltic environmental management in the following 40 years. The loads of many polluting substances have been greatly reduced, but legacy pollution slows recovery. Top predator populations have recovered, and human exposure to potential toxins has been reduced. The cod stock has partially recovered. Nutrient loads are decreasing, but deep-water anoxia and cyanobacterial blooms remain extensive, and climate change threatens the advances made. Ecosystem-based management is the agreed principle, but in practice the various environmental problems are still handled separately, since we still lack both basic ecological knowledge and appropriate governance structures for managing them together, in a true ecosystem approach.

Nitrogen fixation by cyanobacteria stimulates production in Baltic food webs.

Filamentous, nitrogen-fixing cyanobacteria form extensive summer blooms in the Baltic Sea. Their ability to fix dissolved N2 allows cyanobacteria to circumvent the general summer nitrogen limitation, while also generating a supply of novel bioavailable nitrogen for the food web. However, the fate of the nitrogen fixed by cyanobacteria remains unresolved, as does its importance for secondary production in the Baltic Sea. Here, we synthesize recent experimental and field studies providing strong empirical evidence that cyanobacterial nitrogen is efficiently assimilated and transferred in Baltic food webs via two major pathways: directly by grazing on fresh or decaying cyanobacteria and indirectly through the uptake by other phytoplankton and microbes of bioavailable nitrogen exuded from cyanobacterial cells. This information is an essential step toward guiding nutrient management to minimize noxious blooms without overly reducing secondary production, and ultimately most probably fish production in the Baltic Sea.

Baltic Sea ecosystem-based management under climate change: Synthesis and future challenges.

Ecosystem-based management (EBM) has emerged as the generally agreed strategy for managing ecosystems, with humans as integral parts of the managed system. Human activities have substantial effects on marine ecosystems, through overfishing, eutrophication, toxic pollution, habitat destruction, and climate change. It is important to advance the scientific knowledge of the cumulative, integrative, and interacting effects of these diverse activities, to support effective implementation of EBM. Based on contributions to this special issue of AMBIO, we synthesize the scientific findings into four components: pollution and legal frameworks, ecosystem processes, scale-dependent effects, and innovative tools and methods. We conclude with challenges for the future, and identify the next steps needed for successful implementation of EBM in general and specifically for the Baltic Sea.

Nitrogen fixed by cyanobacteria is utilized by deposit-feeders.

Benthic communities below the photic zone depend for food on allochthonous organic matter derived from seasonal phytoplankton blooms. In the Baltic Sea, the spring diatom bloom is considered the most important input of organic matter, whereas the contribution of the summer bloom dominated by diazotrophic cyanobacteria is less understood. The possible increase in cyanobacteria blooms as a consequence of eutrophication and climate change calls for evaluation of cyanobacteria effects on benthic community functioning and productivity. Here, we examine utilization of cyanobacterial nitrogen by deposit-feeding benthic macrofauna following a cyanobacteria bloom at three stations during two consecutive years and link these changes to isotopic niche and variations in body condition (assayed as C:N ratio) of the animals. Since nitrogen-fixing cyanobacteria have δ(15)N close to -2‰, we expected the δ(15)N in the deposit-feeders to decrease after the bloom if their assimilation of cyanobacteria-derived nitrogen was substantial. We also expected the settled cyanobacteria with their associated microheterotrophic community and relatively high nitrogen content to increase the isotopic niche area, trophic diversity and dietary divergence between individuals (estimated as the nearest neighbour distance) in the benthic fauna after the bloom. The three surface-feeding species (Monoporeia affinis, Macoma balthica and Marenzelleria arctia) showed significantly lower δ(15)N values after the bloom, while the sub-surface feeder Pontoporeia femorata did not. The effect of the bloom on isotopic niche varied greatly between stations; populations which increased niche area after the bloom had better body condition than populations with reduced niche, regardless of species. Thus, cyanobacterial nitrogen is efficiently integrated into the benthic food webs in the Baltic, with likely consequences for their functioning, secondary production, transfer efficiency, trophic interactions, and intra- and interspecific competition.

Baltic Sea nutrient reductions--what should we aim for?

Nutrient load reductions are needed to improve the state of the Baltic Sea, but it is still under debate how they should be implemented. In this paper, we use data from an environmental valuation study conducted in all nine Baltic Sea states to investigate public preferences of relevance to three of the involved decision-dimensions: First, the roles of nitrogen versus phosphorus reductions causing different eutrophication effects; second, the role of time - the lag between actions to reduce nutrient loads and perceived improvements; and third; the spatial dimension and the roles of actions targeting the coastal and open sea environment and different sub-basins. Our findings indicate that respondents view and value the Baltic Sea environment as a whole, and are not focussed only on their local sea area, or a particular aspect of water quality. We argue that public preferences concerning these three perspectives should be one of the factors guiding marine policy. This requires considering the entire range of eutrophication effects, in coastal and open sea areas, and including long-term and short-term measures.

Polychaete invader enhances resource utilization in a species-poor system.

Ecosystem consequences of biodiversity change are often studied from a species loss perspective, while the effects of invasive species on ecosystem functions are rarely quantified. In this experimental study, we used isotope tracers to measure the incorporation and burial of carbon and nitrogen from a simulated spring phytoplankton bloom by communities of one to four species of deposit-feeding macrofauna found in the species-poor Baltic Sea. The recently invading polychaete Marenzelleria arctia, which has spread throughout the Baltic Sea, grows more rapidly than the native species Monoporeia affinis, Pontoporeia femorata (both amphipods) and Macoma balthica (a bivalve), resulting in higher biomass increase (biomass production) in treatments including the polychaete. Marenzelleria incorporated and buried bloom material at rates similar to the native species. Multi-species treatments generally had higher isotope incorporation, indicative of utilization of bloom material, than expected from monoculture yields of the respective species. The mechanism behind this observed over-yielding was mainly niche complementarity in utilization of the bloom input, and was more evident in communities including the invader. In contrast, multi-species treatments had generally lower biomass increase than expected. This contrasting pattern suggests that there is little overlap in resource use of freshly deposited bloom material between Marenzelleria and the native species but it is likely that interference competition acts to dampen resulting community biomass. In conclusion, an invasive species can enhance incorporation and burial of organic matter from settled phytoplankton blooms, two processes fundamental for marine productivity.

Diversity of larger consumers enhances interference competition effects on smaller competitors.

Competition between large and small species for the same food is common in a number of ecosystems including aquatic ones. How diversity of larger consumers affects the access of smaller competitors to a limiting resource is not well understood. We tested experimentally how species richness (0-3 spp.) of benthic deposit-feeding macrofauna changes meiofaunal ostracods' incorporation of fresh organic matter from a stable-isotope-labeled cyanobacterial bloom, using fauna from the species-poor Baltic Sea. Presence of macrofauna mostly decreased meiofaunal incorporation of bloom material, depending on the macrofauna species present. As expected, the species identity of macrofauna influenced the incorporation of organic matter by meiofauna. Interestingly, our results show that, in addition, species richness of the macrofauna significantly reduced meiofauna incorporation of freshly settled nitrogen and carbon. With more than one macrofauna species, the reduction was always greater than expected from the single-species treatments. Field data from the Baltic Sea showed a negative correlation between macrofauna diversity and meiofaunal ostracod abundance, as expected from the experimental results. We argue that this is caused by interference competition, due to spatial niche differentiation between macrofauna species reducing the sediment volume in which ostracods can feed undisturbed by larger competitors. Interference from macrofauna significantly reduces organic matter incorporation by meiofauna, indicating that diversity of larger consumers is an important factor controlling the access of smaller competitors to a limiting food resource.

Higher diversity of deposit-feeding macrofauna enhances phytodetritus processing.

The link between biodiversity and ecosystem functioning is an important question that remains unresolved, particularly in marine systems, in which cycling of organic matter by benthic organisms is of global significance. Direct observations of specific resource use by each species in single- and multispecies communities, as quantified by stable isotopes, facilitates a mechanistic understanding of the importance of each species for ecosystem functioning. We tested the effects of altered biodiversity (species richness) of deposit-feeding macrofauna on incorporation and burial of phytodetritus in combinations of three species representing natural communities found in the sediments of the species-poor Baltic Sea. The three species, two amphipods and a bivalve, had different rates of incorporation and burial and different needs for carbon (C) and nitrogen (N). The amphipods exhibited clear resource partitioning in sympatry, as a result of vertical separation in the sediment and consequent differential use of food. Communities of several species incorporated more C and N than expected from the respective single-species treatments, due to higher incorporation by surface feeders in multispecies treatments. Community incorporation of N in the most diverse treatment even exceeded N incorporation by a single-species treatment of the best-performing species, showing transgressive over-yielding. This over-yielding was primarily due to positive complementarity in all treatments. Diverse soft bottoms are also likely to be more productive in the long run, as species-specific traits (subsurface feeding) preserve fresh phytodetritus by burying it to depths in the sediment at which the mineralization rate is low. The more diverse sediment communities showed more efficient trophic transfer of phytodetritus, a finding of general significance for understanding biological processes driving the transformation of nutrients and energy in benthic ecosystems.

Adaptive coastal planning and the European Union's water framework directive: a Swedish perspective.

The implementation of the EU Water Framework Directive and the achievement of its water quality objectives will to a large degree depend on scientific knowledge about the sources and status of water resources and the effects of mitigative measures. Coastal waters will be of central interest, since the management of whole river basins may be affected if coastal water status is not satisfactory. The Water Directive's general directions on what information the water authorities need and on suitable methods for obtaining such information leave it to the Member states to decide on more precise instructions. Improved legal mechanisms are needed to assist and guide the managers and scientists charged with implementing the directive and achieving its water quality objectives, particularly when faced with insufficient information, flawed knowledge, or changed circumstances. We assess progress in the current process of implementing the Water Directive in Sweden, focussing on coastal waters, and on general problems arising using this type of legal technique.

Effects of the deposit-feeding benthic bivalve Macoma balthica on meiobenthos.

To evaluate the effects of a deposit-feeding bivalve on meiobenthic assemblage structure in muddy habitats, a laboratory experiment was performed at the Askö Laboratory in the northwestern Baltic proper. Microcosms, surface area 104 cm2, containing a c. 7-cm thick layer of sieved (0.5 mm) sublittoral mud were established in June 1990. Two months later the tellinid bivalve Macoma balthica was added in quantities varying from 0 to 40 individuals per microcosm. After 5 months the effects of the bivalves on the meiofauna were surprisingly small. The density of harpacticoid copepods was lowest (P <0.05) in microcosms containing a high density of M. balthica. It is suggested that competition for food resources was responsible for this pattern. For all other meiofaunal groups, including nematodes which were the most abundant taxon (99%), no significant differences (P>0.05) were observed among treatments. The assemblage structure of the nematodes was similar between treatments. The vertical distribution of both major taxa and nematode species appeared to be unaffected by the presence of the bivalve.

Predation by the isopod Saduria entomon on the amphipods Monoporeia affinis and Pontoporeia femorata: experiments on prey vulnerability.

Predation by Saduria entomon on the depositfeeding amphipods Monoporeia (=Pontoporeia) affinis and Pontoporeia femorata was studied in laboratory experiments. Prey vulnerabilities were compared in singleprey treatments and in mixed-prey treatments, where the proportions of the two species were varied at one total density. In a pilot experiment, P. femorata was the more vulnerable prey, both in single-prey and mixed-prey treatments. In later experiments, the amphipod species were equally vulnerable in single-prey treatments, while in mixed-prey treatments Saduria preyed preferentially on M. affinis, even when this prey was rare, i.e. it did not "switch". We suggest that the different result in the later experiments may have been due to a change in the search mode of the isopods.