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  • Villnäs, Anna; Janas, Urzsula; Josefson, Alf B.; Kendzierska, Halina; Nygård, Henrik; Norkko, Joanna; Norkko, Alf (2019)
    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.
  • Villnäs, Anna; Norkko, Alf; Lehtonen, Kari K. (2019)
    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.
  • Bernard, Guillaume; Gammal, Johanna; Järnström, Marie; Norkko, Joanna; Norkko, Alf (2019)
    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.