Nutrient fluxes to coastal areas have risen in recent decades, leading to widespread hypoxia and other ecological damage, particularly from nitrogen (N). Several factors make N more limiting in estuaries and coastal waters than in lakes: desorption (release) of phosphorus (P) bound to clay as salinity increases, lack of planktonic N fixation in most coastal ecosystems, and flux of relatively P-rich, N-poor waters from coastal oceans into estuaries. During eutrophication, biogeochemical feedbacks further increase the supply of N and P, but decrease availability of silica - conditions that can favor the formation and persistence of harmful algal blooms. Given sufficient N inputs, estuaries and coastal marine ecosystems can be driven to P limitation. This switch contributes to greater far-field N pollution; that is, the N moves further and contributes to eutrophication at greater distances. The physical oceanography (extent of stratification, residence time, and so forth) of coastal systems determines their sensitivity to hypoxia, and recent changes in physics have made some ecosystems more sensitive to hypoxia. Coastal hypoxia contributes to ocean acidification, which harms calcifying organisms such as mollusks and some crustaceans. (Less)

https://esajournals.onlinelibrary.wiley.com/doi/pdf/10.1890/100008

Coupled biogeochemical cycles: eutrophication and hypoxia in temperate estuaries and coastal marine ecosystems

. Coastal hypoxia (defined here as Hypoxia alters both the structure and function of benthic communities, but effects may differ with regional hypoxia history. Human-caused hypoxia is generally linked to eutrophication, and occurs adjacent to watersheds with large populations or agricultural activities. Many occurrences are seasonal, within estuaries, fjords or enclosed seas of the North Atlantic and the NW Pacific Oceans. Benthic faunal responses, elicited at oxygen levels below 2 ml L−1, typically involve avoidance or mortality of large species and elevated abundances of enrichment opportunists, sometimes prior to population crashes. Areas of low oxygen persist seasonally or continuously beneath upwelling regions, associated with the upper parts of oxygen minimum zones (SE Pacific, W Africa, N Indian Ocean). These have a distribution largely distinct from eutrophic areas and support a resident fauna that is adapted to survive and reproduce at oxygen concentrations

/pdf/effects-of-natural-and-human-induced-hypoxia-on-coastal-2nt30psige.pdf

Effects of natural and human-induced hypoxia on coastal benthos

Eutrophication has been an increasing ecological threat during the past 50 yr in many Scandinavian and Baltic marine waters. Large sedimentary areas are seasonally, or more or less permanently, affected by hypoxia and/or anoxia with devastating effects on the benthic macrofauna in, for example, the Baltic Sea, the Belt Seas and Oresund between Denmark and Sweden, the Kattegat and the Skagerrak coast towards the North Sea. In this review figures for the input of nitrogen and phosphorus to different sea areas are presented, and in several cases also changes of nitrogen and phosphorus concentrations in the water. The nutrient input is related to production levels, and related to macrobenthic infauna. Changes of dominant benthic species, abundance and biomass are presented in relation to both changes in organic enrichment and hypoxia and/or anoxia in time and space. Since the 1950s-60s, the benthic faunal biomass has increased in the Gulf of Bothnia as a result of increased organic enrichment. In the Aland Archipelago, the number of benthic species decreased since the 1970s but abundance and biomass increased. Drifting algae at the sediment surface has also been an increasing problem. The changes were caused by increasing eutrophication. In the Finnish Archipelago Sea, large-scale eutrophication has resulted in periodic bottom water hypoxia and drifting algal mats with negative effects on benthic fauna. In the Gulf of Finland, the benthic fauna has been negatively affected by hypoxic bottom water below 70 in depth since the 1960s, but with a period of improved oxygen conditions during 1987-94. In the Baltic Proper, large sea-bed areas of 70 000-100 000 km(2) below 70-80 in water depth have been more or less hypoxic and/or anoxic since the 1960s with no or reduced sediment-dwelling fauna. This process was a result of increased eutrophication and lack of larger inflows of oxygenated water from the Kattegat. Several coastal areas and larger basins in the southern Baltic (e.g. the Bornholm Basin, the Arkona Basin and the Kiel Bay), have, on occasions, been similarly negatively affected by hypoxic bottom water. Many sedimentary areas below similar to 17 in in the Danish Belt Seas have been affected by seasonal hypoxia since the 1970s with negative consequences for the bottom fauna. On the Danish Kattegat coast, the benthic fauna in the Limfjord, the Mariager fjord and the Roskilde fjord have been particularly negatively affected. In the southeast, open Kattegat, increased input of nutrients in combination with stratification have resulted in seasonal hypoxia since 1980 with negative effects on benthic animals and commercial fish species in most years. Several fjords on the Swedish and Norwegian Skagerrak coast have shown negative temporal trends in bottom water oxygen concentrations, and some of them lack benthic fauna in the deeper parts for several months or more. In this review the temporal development of bottom water hypoxia and/or anoxia is discussed and consequent possible losses of sediment-dwelling faunal biomass are roughly calculated. In total for the areas investigated, the worst years of hypoxia and/or anoxia combined may have reduced the benthic macrofaunal biomass by 3 million t. This loss is partly compensated by the biomass increase that has occurred in well-flushed organically enriched coastal areas. Tolerance of some Baltic species to hypoxia and/or anoxia is discussed and also their different strategies to cope with hypoxia and/or anoxia and H2S.

Temporal and spatial large-scale effects of eutrophication and oxygen deficiency on benthic fauna in Scandinavian and Baltic waters - A review

In aquatic ecosystems, invertebrate bioturbation significantly influences microbial activities and biogeochemical processes in sediments by modifying water and sediment fluxes at the water-sediment interface. We apply the concept of ecosystem engineering to develop a qualitative general understanding of the role of bioturbation on microbial processes in different benthic environments. We hypothesized that the effects of the bioturbation mode (sediment reworking, biogenic structure building, bioirrigation) on microbial processes vary between diffusion- and advection-dominated benthic environments because bioturbation does not have the same influences on hydrological exchanges (and the flux of resources for micro-organisms living in sediments) at the water-sediment interface of the two systems. To test this hypothesis, we experimentally compared the influence of three bioturbation modes (fine-sediment reworking, U-shaped structure burrowing, and gallery-network burrowing) in a diffusion-dominated system (fine sediments/low interstitial flow rates) and an advectiondominated system (coarse sediments/advection of water in sediments). Our analysis demonstrated that bioturbation modes in the two systems had different impacts on microbial activities. For instance, U-shaped tube burrowing by animals increased O2 consumption in the diffusion- dominated system but produced the opposite effect in the advection-dominated system. The influence of bioturbation was also negatively related to interstitial flow rate, the bioturbation having a higher influence on O2 consumption in the diffusion-dominated system than in the advection-dominated system. According to our hypothesis, bioturbation modified microbial processes in sediments depend on the hydrological characteristics of the system. In the diffusion-dominated system, invertebrate bioturbation can produce water fluxes at the watersediment interface that may strongly influence microbial processes in sediments. In contrast, in the advectiondominated system, invertebrate bioturbation can only modify the water circulation patterns in sediments, moderately affecting microbial processes. Consequently, it is necessary to use a conceptual framework which takes into account the features of sediment habitats in order to allow a better prediction of bioturbation effects on sediment biogeochemistry. With this aim, the conceptual scheme of ecosystem engineers can be an organizing principle to integrate the complex relationships among physical habitat, bioturbation mode and microbial activity.

Ecosystem engineering: the impact of bioturbation on biogeochemical processes in marine and freshwater benthic habitats

Benthic invertebrates play a key role in the physical, chemical, and biological properties of the marine water-sediment interface. The influences of invertebrates on biogeochemical processes have mainly been attributed to their sediment reworking and bioirrigation activities. The aim of this study was to compare the influences of bioturbation activities by 3 dominant species of shallow water habitats (Cerastoderma edule, Corophium volutator, and Nereis diversicolor) on microbial communi- ties and biogeochemical processes in sediment cores. C. edule acted as a biodiffuser, mixing surface particles in the top 2 cm of the sediment. Despite this mixing activity, this species had little effect on O2 consumption, water exchange between the water column and the sediment, microbial character- istics, and release of nutrients from the sediment. In contrast, C. volutator and N. diversicolor produced burrows in the sediment that allowed transport of surface particles into biogenic structures. These 2 species doubled the solute exchange between the water column and the sediment. Such modifications of sediment structure and solute transport increased the O2 consumption and the release of nutrients from the sediment. Both C. volutator and N. diversicolor stimulated the microbial communities as indicated by higher percentages of active bacteria. Reduction of the numbers of sulphate reducing bacteria was observed when the 3 invertebrates were present and could be attrib- uted to the penetration of O2 due to animal activities. N. diversicolor had a greater influence than C. volutator on pore water chemistry, ammonium release, and active bacteria. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediment. The modes of sediment reworking and structure building, irrigation behaviour, and burrowing depths were factors sufficient to assign the 3 species into different functional groups.

https://www.int-res.com/articles/ame2004/36/a036p271.pdf

Influence of bioturbation by three benthic infaunal species on microbial communities and biogeochemical processes in marine sediment

This paper is a brief review of successional stages and activity of benthic soft-bottom communities. Benthic communities was first described by Petersen in the 1910s and further developed by Molander, Thorson and Margalef. Successional stages of benthic communities chance in a predictable way in relation to environmental disturbance and food availability. Food supply to the bottom can occur as a vertical flux, but transport through lateral advection is more important in some areas. While at the bottom, the infauna processes the food in many different ways, and the feeding modes can be categorised into more than 20 functional groups, but fewer are present in brackish water. This categorisation is based on animal mobility and where and how they ingest the food. Animal activity in the sediment, bioturbation, has a significant effect on redox conditions and diagenetic processes. Structures in the sediment due to infaunal presence and activity can be observed in situ by sediment profile imaging, and the biogenic structures and redox conditions can be parameterised and have been shown to correlate to benthic community successional stages. The largest threat to benthic faunal biodiversity is the spread of near-bottom oxygen deficiency in many enclosed are stratified coastal areas.

/pdf/marine-benthic-faunal-successional-stages-and-related-2yrkewdoj6.pdf

Rutger Rosenberg

Papers

Marine benthic faunal successional stages and related sedimentary activity