Abstract. Coastal hypoxia (defined here as <1.42 ml L −1 ; 62.5 µM; 2 mg L −1 , approx. 30% oxygen saturation) develops seasonally in many estuaries, fjords, and along open coasts as a result of natural upwelling or from anthropogenic eutrophication induced by riverine nutrient inputs. Permanent hypoxia occurs naturally in some isolated seas and marine basins as well as in open slope oxygen minimum zones. Responses of benthos to hypoxia depend on the duration, predictability, and intensity of oxygen depletion and on whether H 2 S is formed. Under suboxic conditions, large mats of filamentous sulfide oxidizing bacteria cover the seabed and consume sulfide. They are hypothesized to provide a detoxified microhabitat for eukaryotic benthic communities. Calcareous foraminiferans and nematodes are particularly tolerant of low oxygen concentrations and may attain high densities and dominance, often in association with microbial mats. When oxygen is sufficient to support metazoans, small, soft-bodied invertebrates (typically annelids), often with short generation times and elaborate branchial structures, predominate. Large taxa are more sensitive than small taxa to hypoxia. Crustaceans and echinoderms are typically more sensitive to hypoxia, with lower oxygen thresholds, than annelids, sipunculans, molluscs and cnidarians.

Abstract. Dissolved oxygen (DO) concentration in the water column is an environmental parameter that is crucial for the successful development of many pelagic organisms. Hypoxia tolerance and threshold values are species-and stagespecific and can vary enormously. While some fish species may suffer from oxygen values of less than 3 mL O 2 L −1 through impacted growth, development and behaviour, other organisms such as euphausiids may survive DO levels as low as 0.1 mL O 2 L −1 . A change in the average or the range of DO may have significant impacts on the survival of certain species and hence on the species composition in the ecosystem with consequent changes in trophic pathways and productivity.Evidence for the deleterious effects of oxygen depletion on pelagic species is scarce, particularly in terms of the effect of low oxygen on development, recruitment and patterns of migration and distribution. While planktonic organisms have to cope with variable DOs and exploit adaptive mechanisms, nektonic species may avoid areas of unfavourable DO and develop adapted migration strategies. Planktonic organisms may only be able to escape vertically, above or beneath the Oxygen Minimum Zone (OMZ). In shallow areas only the surface layer can serve as a refuge, but in deep waters many organisms have developed vertical migration strategies to use, pass through and cope with the OMZ.Correspondence to: W. Ekau (wekau@zmt-bremen.de) This paper elucidates the role of DO for different taxa in the pelagic realm and the consequences of low oxygen for foodweb structure and system productivity. We describe processes in two contrasting systems, the semi-enclosed Baltic Sea and the coastal upwelling system of the Benguela Current to demonstrate the consequences of increasing hypoxia on ecosystem functioning and services.

Abstract. Hypoxia has become a world-wide phenomenon in the global coastal ocean and causes a deterioration of the structure and function of ecosystems. Based on the collective contributions of members of SCOR Working Group #128, the present study provides an overview of the major aspects of coastal hypoxia in different biogeochemical provinces, including estuaries, coastal waters, upwelling areas, fjords and semi-enclosed basins, with various external forcings, ecosysCorrespondence to: J. Zhang (jzhang@sklec.ecnu.edu.cn) tem responses, feedbacks and potential impact on the sustainability of the fishery and economics. The obvious external forcings include freshwater runoff and other factors contributing to stratification, organic matter and nutrient loadings, as well as exchange between coastal and open ocean water masses. Their different interactions set up mechanisms that drive the system towards hypoxia. Coastal systems also vary in their relative susceptibility to hypoxia depending on their physical and geographic settings. It is understood that coastal hypoxia has a profound impact on the sustainability of ecosystems, which can be seen, for example, by the change in the food-web structure and system function; other Published by Copernicus Publications on behalf of the European Geosciences Union. 1444 J. Zhang et al.: Natural and human-induced hypoxia and consequences for coastal areas influences include compression and loss of habitat, as well as changes in organism life cycles and reproduction. In most cases, the ecosystem responds to the low dissolved oxygen in non-linear ways with pronounced feedbacks to other compartments of the Earth System, including those that affect human society. Our knowledge and previous experiences illustrate that there is a need to develop new observational tools and models to support integrated research of biogeochemical dynamics and ecosystem behavior that will improve confidence in remediation management strategies for coastal hypoxia.

Abstract. Coastal hypoxia (<1.42 ml L−1; 62.5 μM; 2 mg L−1, approx. 30% oxygen saturation) occurs seasonally in many estuaries, fjords, and along open coasts subject to upwelling or excessive riverine nutrient input, and permanently in some isolated seas and marine basins. Underlying causes of hypoxia include enhanced nutrient input from natural causes (upwelling) or anthropogenic origin (eutrophication) and reduction of mixing by limited circulation or enhanced stratification; combined these lead to higher surface water production, microbial respiration and eventual oxygen depletion. Advective inputs of low-oxygen waters may initiate or expand hypoxic conditions. Responses of estuarine, enclosed sea, and open shelf benthos to hypoxia depend on the duration, predictability, and intensity of oxygen depletion and on whether H2S is formed. Under suboxic conditions, large mats of filamentous sulfide oxidizing bacteria cover the seabed and consume sulfide, thereby providing a detoxified microhabitat for eukaryotic benthic communities. Calcareous foraminiferans and nematodes are particularly tolerant of low oxygen concentrations and may attain high densities and dominance, often in association with microbial mats. When oxygen is sufficient to support metazoans, small, soft-bodied invertebrates (typically annelids), often with short generation times and elaborate branchial structures, predominate. Large taxa are more sensitive than small taxa to hypoxia. Crustaceans and echinoderms are typically more sensitive to hypoxia, with lower oxygen thresholds, than annelids, sipunculans, molluscs and cnidarians. Mobile fish and shellfish will migrate away from low-oxygen areas. Within a species, early life stages may be more subject to oxygen stress than older life stages. 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 <0.5 ml L−1. Under both natural and eutrophication-caused hypoxia there is loss of diversity, through attrition of intolerant species and elevated dominance, as well as reductions in body size. These shifts in species composition and diversity yield altered trophic structure, energy flow pathways, and corresponding ecosystem services such as production, organic matter cycling and organic C burial. Increasingly the influences of nature and humans interact to generate or exacerbate hypoxia. A warmer ocean is more stratified, holds less oxygen, and may experience greater advection of oxygen-poor source waters, making new regions subject to hypoxia. Future understanding of benthic responses to hypoxia must be established in the context of global climate change and other human influences such as overfishing, pollution, disease, habitat loss, and species invasions.