Ninety years of coastal monitoring reveals baseline and extreme ocean temperatures are increasing off the Finnish coast

Long term coastal surface and sub-surface monitoring reveals that warm temperature extremes have intensified and cold extremes nearly vanished in the last decades, increasing baseline water temperature on the southwest coast of Finland. Global marine heatwave assessments often rely on satellite-derived sea surface temperature. However, these data have low accuracy in coastal areas, are unable to measure sub-surface temperatures and have only been available since the 1980s. Here, we analyse 90 years of in situ surface and bottom (30 m) water temperature data from a Finnish coastal monitoring site. Water temperatures were significantly higher between 1991-2020 than 1931-1960 and 1961-1990. We find strong differences between satellite-derived and in situ temperatures, with in situ temperatures being lower in autumn and winter and higher in spring. Measurements at the seafloor indicate marine heatwaves occurred during all seasons between 2016 and 2020, with intensities and durations exceeding previous records. Since the 1990s, we find an upward shift of the baseline temperature and increasingly frequent occurrence of temperatures previously considered as an extreme. Our findings highlight the importance of long-term in situ data and choice of climatological reference periods for assessing change.Peer reviewe

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.Peer reviewe

Seasonal variability in ecosystem functions: quantifying the contribution of invasive species to nutrient cycling in coastal ecosystems

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Increasing densities of an invasive polychaete enhance bioturbation with variable effects on solute fluxes

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.Peer reviewe

Changes in macrofaunal biological traits across estuarine gradients : implications for the coastal nutrient filter

Benthic macrofaunal communities have a profound impact on organic matter turnover and nutrient cycling in marine sediments. Their activities are of particular importance in the coastal filter, where nutrients and organic matter from land are transformed and/or retained before reaching the open sea. The benthic fauna modify the coastal filter directly (through consumption, respiration, excretion and biomass production) and indirectly (through bioturbation). It is hard to experimentally quantify faunal contribution to the coastal filter over large spatial and temporal scales that encompass significant environmental and biological heterogeneity. However, estimates can be obtained with biological trait analyses. By using benthic biological traits, we explored how the potential contribution of macrofaunal communities to the coastal filter differ between inner and outer sites in an extensive archipelago area and examine the generality of the observed pattern across contrasting coastal areas of the entire Baltic Sea. Estimates of benthic bioturbation, longevity and size (i.e. ‘stability’) and total energy and nutrient contents differed between coastal areas and inner versus outer sites. Benthic traits indicative of an enhanced nutrient turnover but a decreased capacity for temporal nutrient retention dominated inner sites, while outer sites were often dominated by larger individuals, exhibiting traits that are likely to enhance nutrient uptake and retention. The overarching similarities in benthic trait expression between more eutrophied inner vs. less affected outer coastal sites across the Baltic Sea suggest that benthic communities might contribute in a similar manner to nutrient recycling and retention in the coastal filter over large geographical scales.peerReviewe

Quantifying bioturbation across coastal seascapes : Habitat characteristics modify effects of macrofaunal communities

Bioturbation by benthic macrofauna communities plays a significant role in the setting and maintenance of important ecosystem functions and the delivery of associated ecosystem services. We investigated the context-dependence of bioturbation performed by natural benthic communities in the coastal northern Baltic Sea by quantifying three bioturbation metrics (particle mixing intensity, surface sediment reworking and bioturbation depth) across 18 sites ranging from cohesive muddy sediments to non-cohesive coarse sands, while accounting for the complexity of natural communities and habitat characteristics. We identified two distinct patterns of bioturbation; in fine sediments bioturbation rates were highly variable and in coarse sediments bioturbation rates were less variable and characterized by lower maximal values. Using distance-based linear multiple regressions, we found that 75.5% of the variance in bioturbation rates in fine sediment could be explained by key functional groups/species abundance and/or biomass (i.e. biomass of the gallery-diffusors and abundances of biodiffusors, surface modifiers, conveyors and gallery diffusors, respectively). In coarse sediment, 47.8% of the variance in bioturbation rates could be explained by a combination of environmental factors (grain size, organic matter content, buried plant material) and faunal functional groups, although fauna alone explained only 13% of this variance. Bioturbation in fine sediments was therefore more predictable based on the composition of benthic fauna. In coarse sediment, the bioturbation activities of benthic fauna were strongly modified by habitat characteristics (including the presence of buried plant material, sediment organic content and grain size) whereas in fine sediments this was not the case. Our results therefore highlight that variability in spatial patterns of bioturbation is a result of complex relationships between macrofauna community structure, sediment type and other habitat characteristics, likely modifying bioturbation performance of individual fauna.Peer reviewe

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.Peer reviewe

Stable seasonal and annual alpha diversity of benthic diatom communities despite changing community composition

The global biodiversity loss has raised interest in the different facets of diversity, and the importance of diversity for ecosystem functions has been recognized. However, our knowledge on seasonal and inter-annual variation in the composition and diversity of communities is still poor. Here, we investigated the seasonal and inter-annual changes in taxonomic and functional community composition and diversity of benthic diatoms in a coastal habitat of the northern Baltic Sea, where seasonal and inter-annual variation of climate is pronounced. We found that the taxonomic and functional alpha diversity remained stable at seasonal and inter-annual level despite strong changes in community composition. However, alpha diversity decreased during an exceptionally warm winter possibly due to disturbances induced by the lack of ice. This may suggest that climate warming and consequently limited ice cover will affect the diversity of benthic communities.Peer reviewe

Species and functional trait turnover in response to broad-scale change and an invasive species

While beta diversity has been implicated as a key factor in controlling resilience of communities to stressors, lack of long-term data sets has limited the study of temporal dynamics of beta diversity. With a time series at two sites in excess of 40yr, we investigated turnover of both species and functional traits in a system stressed by eutrophication and overfishing and undergoing climate change and invasion. The two sites, although located near to each other, differ in water depth (20 cf. 35m), but both sites have displayed increased abundances of an invasive polychaete since 1990. We tested two hypotheses related to the effect of an invasive species; that taxa richness and turnover would decrease, and trait richness would increase post invasion and that trait turnover would increase between arrival and establishment of the invasive. Generally, we observed different dynamics at the two sites and responses not consistent with our hypotheses. We detected an increase in taxa richness at both sites and an increase in taxa turnover and number of traits at one site only. Trait turnover was higher prior to the invasion, although again only at one site. Disjunctive responses between species and trait turnover occurred, with the invader contributing in a nonrandom fashion to trait turnover. The lack of strong, consistent responses to the arrival and establishment of the invasive, and the decrease in trait turnover, suggests that effects of invasives are not only system- and species-dependent, but also depend on community dynamics of the invaded site, in particular the assembly processes, and historical context.Peer reviewe

Diversity and distribution across a large environmental and spatial gradient: Evaluating the taxonomic and functional turnover, transitions and environmental drivers of benthic diatom communities

Aim Global biodiversity loss has raised interest in understanding variation in diversity at different scales. In particular, studies conducted across large spatial gradients are crucial, because they can increase perspectives on how ecological patterns change relative to environmental factors and facilitate predictions of possible responses to environmental change. We explored the full extent of a brackish sea to test the hypotheses that: (a) benthic communities are defined by the limited ranges of species, controlled by varying drivers along a large environmental gradient; (b) the responses of taxonomic and functional community composition and turnover to the environmental gradient are different, thus highlighting the need to include both measures in ecological studies; and (c) diversity reaches the minimum at intermediate salinities (Remane curve) owing to the low adaptation of freshwater and marine species. Location A large environmental and spatial gradient spanning the entire Swedish coastline (c. 2,300 km; salinity 1.2-27.6), the Baltic Sea. Time period August 2018. Major taxa studied Benthic diatoms. Methods We assessed environmental drivers for the communities and calculated the taxonomic and functional alpha and beta diversity along the gradient. We also compared the taxonomic and functional composition and diversity of communities among areas with different salinity. Results We found support for the hypothesis of limited species ranges, because taxonomic beta diversity, mainly induced by changes in salinity and climate, was high, whereas functional beta diversity remained considerably lower, and the composition and diversity of communities, in addition to environmental drivers controlling the communities, differed between regions with different salinity. The lowest taxonomic diversity was found at intermediate salinities of 5-6. Main conclusions These findings advance understanding of large-scale patterns of benthic diversity, emphasize the importance of large gradient studies for a better understanding of general ecological patterns and highlight the vulnerability of brackish water ecosystems as ecologically important tipping-point realms.Peer reviewe