Seasonal population dynamics of the invasive polychaete genus Marenzelleria spp. in contrasting soft-sediment habitats

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.Peer reviewe

Estimating respiration rates and secondary production of macrobenthic communities across coastal habitats with contrasting structural biodiversity

A central goal of benthic ecology is to describe the pathways and quantities of energy and material flow in seafloor communities over different spatial and temporal scales. We examined the relative macrobenthic contribution to the seafloor metabolism by estimating respiration and secondary production based on seasonal measurements of macrofauna biomass across key coastal habitats of the Baltic Sea archipelago. Then, we compared the macrofauna estimates with estimates of overall seafloor gross primary production and respiration obtained from the same habitats using the aquatic eddy covariance technique. Estimates of macrobenthic respiration rates suggest habitat-specific macrofauna contribution (%) to the overall seafloor respiration ranked as follows: blue mussel reef (44.5) > seagrass meadow (25.6) > mixed meadow (24.1) > bare sand (17.8) > Fucus-bed (11.1). In terms of secondary production (g C m−2 y−1), our estimates suggest ranking of habitat value as follows: blue mussel reef (493.4) > seagrass meadow (278.5) > Fucus-bed (102.2) > mixed meadow (94.2) > bare sand (52.1). Our results suggest that approximately 12 and 10% of the overall soft-sediment metabolism translated into macrofauna respiration and secondary production, respectively. The hard-bottoms exemplified two end-points of the coastal metabolism, with the Fucus-bed as a high producer and active exporter of organic C (that is, net autotrophy), and the mussel reef as a high consumer and active recycler of organic C (that is, net heterotrophy). Using a combination of metrics of ecosystem functioning, such as respiration rates and secondary production, in combination with direct habitat-scale measurements of O2 fluxes, our study provides a quantitative assessment of the role of macrofauna for ecosystem functioning across heterogeneous coastal seascapes.Peer reviewe

Temporal Variation in Resuspension Potential and Associated Nutrient Dynamics in Shallow Coastal Environments

Sediment resuspension may play a major role in sediment-water exchange of nutrients, matter and energy in coastal areas where waves and currents dominate sediment transport. Biogeochemical sediment properties regulate sediment erodibility, but there is only limited knowledge of how temporal variability in environmental variables is reflected in the resuspension potential, especially for subtidal habitats. Further, the significance of resuspension on nutrient fluxes in coastal environments has remained unclear as contradicting results have been reported. Here we quantified the temporal variation in resuspension potential metrics (erosion threshold (τc; N m−2) and erosion constant (me; g N−1 s−1)) and associated nutrient fluxes from three sites in the Hanko archipelago (Finland) using a core-based erosion device (EROMES). The sites were sampled bi-monthly from April to December. We also quantified the temporal variation in biogeochemical sediment properties at each site. The τc exhibited the clearest temporal pattern in muddy sediment, where the coefficient of variation (= 67) was two to three times higher than the mixed (= 29) and sandy (= 16) sediments. Dry bulk density was the best predictor for sediment erodibility at all sites explaining 26–46% of the temporal variation in τc despite its limited variability at sandier sites. In addition, temporal variations in the macrofaunal community were important predictors of muddy sediment erodibility and therefore community dynamics need to be considered in sediment transport studies. All sites were potential nutrient sources, yet the overall role of sediment resuspension on nutrient release from the sediments was small.Peer reviewe

Coastal hypoxia and the importance of benthic macrofauna communities for ecosystem functioning

Coastal ecosystems are important because of the vital ecosystem functions and services they provide, but many are threatened by eutrophication and hypoxia. This results in loss of biodiversity and subsequent changes in ecosystem functioning. Consequently, the need for empirical field studies regarding biodiversity-ecosystem functioning in coastal areas has been emphasized. The present field study quantified the links between benthic macrofaunal communities (abundance, biomass, and species richness), sediment oxygen consumption, and solute fluxes (NO3- + NO2-, NH4+, PO43-, SiO4, Fe, Mn) along a 7.5-km natural gradient of seasonal hypoxia in the coastal northern Baltic Sea. Sampling was done in late August 2010 in the middle archipelago zone of the Hanko peninsula, Finland. As predicted, the macrofaunal communities were decimated with increasing hypoxia, and the nutrient transformation processes were changed at the sediment-water interface, with notably higher effluxes of phosphate and ammonium from the sediment. Solute fluxes varied even during normoxia, which implies a high context-dependency, and could be explained by even small variations in environmental variables such as organic matter and C/N ratios. Importantly, the low diversity benthic macrofaunal communities, which were dominated by Macoma balthica and the invasive Marenzelleria spp., had a large influence on the solute fluxes, especially under normoxia, but also under hypoxia.Peer reviewe

Characterization of soft-bottom benthic habitats of the Aland islands, northern Baltic sea

Sediment surface and profile imaging (SPI) was used in combination with grab sampling of sediment (sediment type, organic content, benthic infauna) and hydrography (temperature, oxygen saturation of bottom water) to analyze and describe the soft-bottom benthic habitats of the Aland archipelago (60 degrees 00\u27 to 60 degrees 30\u27 N, 19 degrees 30\u27 to 20 degrees 30\u27 E) in the northern Baltic Sea. The SPI analysis covered 42 stations (5 to 263 m depth), from inner sheltered bays to open coastal waters, with varying sediment types (soft mud with high organic content to sandy substrates with low organic content; loss on ignition: 0.5 to 12.4%). Clustering of the sampled stations (sediment properties) yielded 3 distinct categories of sedimentary habitats: (1) inner archipelago areas and bays with high organic content of the sediment and reduced oxygen saturation in the bottom water, (2) archipelago waters with intermediate values of all analyzed parameters, and (3) open coastal sediments with low organic content and high oxygen saturation (2 deep offshore stations formed an additional group based primarily on depth). Visual analysis of the images provided information on several additional abiotic and biotic characteristics of the sediment, and significant correlations were found mainly between oxygen saturation, organic content, sediment type, shear strength (penetration of gear), surface relief and the depth of the redox potential discontinuity layer in the sediment. The sediment properties were also reflected in the zoobenthos. The correlations between parameters measured are discussed in relation to applicability of the SPI method, monitoring demands, and basic understanding of the sediment-animal relationships

Identifying areas prone to coastal hypoxia - the role of topography

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.Peer reviewe

Intermittent bioirrigation and oxygen dynamics in permeable sediments: An experimental and modeling study of three tellinid bivalves

To explore the dynamic nature of geochemical conditions in bioirrigated marine permeable sediments, we studied the hydraulic activity of three tellinacean bivalve molluscs (the Pacific species Macoma nasuta and Macomona liliana, and the northern Atlantic and Pacific species Macoma balthica). We combined porewater pressure sensing, time-lapse photography and oxygen imaging to quantify the durations and frequencies of tellinid irrigation activity and the associated oxygen dynamics in the sediment. Porewater pressure records of all tellinids were dominated by intermittent porewater pressurization, induced by periodic water injection into the sediment through their excurrent siphons, which resulted in intermittent oxygen supply to subsurface sediments. The irrigation (two–12 minutes long) and intervals between subsequent irrigation bouts (1.5–13 minutes) varied among tellinid species and individual sizes. For large M. liliana and M. nasuta, the average intervals between irrigation bouts were sufficiently long (10 minutes and four minutes, respectively) to allow complete oxygen consumption in between irrigation bouts in all tested sediment types. Irrigation patterns of smaller conspecifics and the smaller species M. balthica were characterized by significantly shorter separation of irrigation bouts, which resulted in more continuous oxygenation of the sediment. Transport-reaction modeling confirmed these species- and size-specific geochemical signatures and indicated that the geochemical character of the sediment is largely conditioned by the interplay between temporal irrigation patterns and sedimentary oxygen consumption rates. For large tellinids, model simulations indicated that oscillatory rather than stationary geochemical conditions are prevalent in a wide range of sediment types, with oxic pockets collapsing completely between periods of active irrigation. Based on the model results we developed analytical approximations that allow estimation of spatio-temporal characteristics of sediment oxygenation for a wide range of sediment types and infaunal activity patterns. Our results emphasize the need to consider the intermittent nature of bioirrigation when studying the geochemical impact of infauna in permeable sediments

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.Peer reviewe

High throughput shotgun sequencing of eRNA reveals taxonomic and derived functional shifts across a benthic productivity gradient

Benthic macrofauna is regularly used in monitoring programmes, however the vast majority of benthic eukaryotic biodiversity lies mostly in microscopic organisms, such as meiofauna (invertebrates <1 mm) and protists, that rapidly responds to environmental change. These communities have traditionally been hard to sample and handle in the laboratory, but DNA sequencing has made such work less time consuming. While DNA sequencing captures both alive and dead organisms, environmental RNA (eRNA) better targets living organisms or organisms of recent origin in the environment. Here, we assessed the biodiversity of three known bioindicator microeukaryote groups (nematodes, foraminifera, and ciliates) in sediment samples collected at seven coastal sites along an organic carbon (OC) gradient. We aimed to investigate if eRNA shotgun sequencing can be used to simultaneously detect differences in (i) biodiversity of multiple microeukaryotic communities; and (ii) functional feeding traits of nematodes. Results showed that biodiversity was lower for nematodes and foraminifera in high OC (6.2%-6.9%), when compared to low OC sediments (1.2%-2.8%). Dissimilarity in community composition increased for all three groups between Low OC and High OC, as well as the classified feeding type of nematode genera (with more nonselective deposit feeders in high OC sediment). High relative abundant genera included nematodeSabatieriaand foraminiferaElphidiumin high OC, andCryptocaryon-like ciliates in low OC sediments. Considering that future sequencing technologies are likely to decrease in cost, the use of eRNA shotgun sequencing to assess biodiversity of benthic microeukaryotes could be a powerful tool in recurring monitoring programmes.Peer reviewe

Sediment properties, biota and local habitat structure explain variation in the erodibility of coastal sediments

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.Peer reviewe