. Water masses can become undersaturated with oxygen when natural processes alone or in combination with anthropogenic processes produce enough organic carbon that is aerobically decomposed faster than the rate of oxygen re-aeration. The dominant natural processes usually involved are photosynthetic carbon production and microbial respiration. The re-supply rate is indirectly related to its isolation from the surface layer. Hypoxic water masses ( Hypoxia development and continuation in many areas of the world's coastal ocean is accelerated by human activities, especially where nutrient loading increased in the Anthropocene. This higher loading set in motion a cascading set of events related to eutrophication. The formation of hypoxic areas has been exacerbated by any combination of interactions that increase primary production and accumulation of organic carbon leading to increased respiratory demand for oxygen below a seasonal or permanent pycnocline. Nutrient loading is likely to increase further as population growth and resource intensification rises, especially with increased dependency on crops using fertilizers, burning of fossil fuels, urbanization, and waste water generation. It is likely that the occurrence and persistence of hypoxia will be even more widespread and have more impacts than presently observed. Global climate change will further complicate the causative factors in both natural and human-caused hypoxia. The likelihood of strengthened stratification alone, from increased surface water temperature as the global climate warms, is sufficient to worsen hypoxia where it currently exists and facilitate its formation in additional waters. Increased precipitation that increases freshwater discharge and flux of nutrients will result in increased primary production in the receiving waters up to a point. The interplay of increased nutrients and stratification where they occur will aggravate and accelerate hypoxia. Changes in wind fields may expand oxygen minimum zones onto more continental shelf areas. On the other hand, not all regions will experience increased precipitation, some oceanic water temperatures may decrease as currents shift, and frequency and severity of tropical storms may increase and temporarily disrupt hypoxia more often. The consequences of global warming and climate change are effectively uncontrollable at least in the near term. On the other hand, the consequences of eutrophication-induced hypoxia can be reversed if long-term, broad-scale, and persistent efforts to reduce substantial nutrient loads are developed and implemented. In the face of globally expanding hypoxia, there is a need for water and resource managers to act now to reduce nutrient loads to maintain, at least, the current status.

/pdf/dynamics-and-distribution-of-natural-and-human-caused-21e6uqwqxc.pdf

Dynamics and distribution of natural and human-caused hypoxia

Oceanography and marine biology: an annual review

. The deep sea, the largest biome on Earth, has a series of characteristics that make this environment both distinct from other marine and land ecosystems and unique for the entire planet. This review describes these patterns and processes, from geological settings to biological processes, biodiversity and biogeographical patterns. It concludes with a brief discussion of current threats from anthropogenic activities to deep-sea habitats and their fauna. Investigations of deep-sea habitats and their fauna began in the late 19th century. In the intervening years, technological developments and stimulating discoveries have promoted deep-sea research and changed our way of understanding life on the planet. Nevertheless, the deep sea is still mostly unknown and current discovery rates of both habitats and species remain high. The geological, physical and geochemical settings of the deep-sea floor and the water column form a series of different habitats with unique characteristics that support specific faunal communities. Since 1840, 28 new habitats/ecosystems have been discovered from the shelf break to the deep trenches and discoveries of new habitats are still happening in the early 21st century. However, for most of these habitats the global area covered is unknown or has been only very roughly estimated; an even smaller – indeed, minimal – proportion has actually been sampled and investigated. We currently perceive most of the deep-sea ecosystems as heterotrophic, depending ultimately on the flux on organic matter produced in the overlying surface ocean through photosynthesis. The resulting strong food limitation thus shapes deep-sea biota and communities, with exceptions only in reducing ecosystems such as inter alia hydrothermal vents or cold seeps. Here, chemoautolithotrophic bacteria play the role of primary producers fuelled by chemical energy sources rather than sunlight. Other ecosystems, such as seamounts, canyons or cold-water corals have an increased productivity through specific physical processes, such as topographic modification of currents and enhanced transport of particles and detrital matter. Because of its unique abiotic attributes, the deep sea hosts a specialized fauna. Although there are no phyla unique to deep waters, at lower taxonomic levels the composition of the fauna is distinct from that found in the upper ocean. Amongst other characteristic patterns, deep-sea species may exhibit either gigantism or dwarfism, related to the decrease in food availability with depth. Food limitation on the seafloor and water column is also reflected in the trophic structure of heterotrophic deep-sea communities, which are adapted to low energy availability. In most of these heterotrophic habitats, the dominant megafauna is composed of detritivores, while filter feeders are abundant in habitats with hard substrata (e.g. mid-ocean ridges, seamounts, canyon walls and coral reefs). Chemoautotrophy through symbiotic relationships is dominant in reducing habitats. Deep-sea biodiversity is among of the highest on the planet, mainly composed of macro and meiofauna, with high evenness. This is true for most of the continental margins and abyssal plains with hot spots of diversity such as seamounts or cold-water corals. However, in some ecosystems with particularly "extreme" physicochemical processes (e.g. hydrothermal vents), biodiversity is low but abundance and biomass are high and the communities are dominated by a few species. Two large-scale diversity patterns have been discussed for deep-sea benthic communities. First, a unimodal relationship between diversity and depth is observed, with a peak at intermediate depths (2000–3000 m), although this is not universal and particular abiotic processes can modify the trend. Secondly, a poleward trend of decreasing diversity has been discussed, but this remains controversial and studies with larger and more robust data sets are needed. Because of the paucity in our knowledge of habitat coverage and species composition, biogeographic studies are mostly based on regional data or on specific taxonomic groups. Recently, global biogeographic provinces for the pelagic and benthic deep ocean have been described, using environmental and, where data were available, taxonomic information. This classification described 30 pelagic provinces and 38 benthic provinces divided into 4 depth ranges, as well as 10 hydrothermal vent provinces. One of the major issues faced by deep-sea biodiversity and biogeographical studies is related to the high number of species new to science that are collected regularly, together with the slow description rates for these new species. Taxonomic coordination at the global scale is particularly difficult, but is essential if we are to analyse large diversity and biogeographic trends. Because of their remoteness, anthropogenic impacts on deep-sea ecosystems have not been addressed very thoroughly until recently. The depletion of biological and mineral resources on land and in shallow waters, coupled with technological developments, are promoting the increased interest in services provided by deep-water resources. Although often largely unknown, evidence for the effects of human activities in deep-water ecosystems – such as deep-sea mining, hydrocarbon exploration and exploitation, fishing, dumping and littering – is already accumulating. Because of our limited knowledge of deep-sea biodiversity and ecosystem functioning and because of the specific life-history adaptations of many deep-sea species (e.g. slow growth and delayed maturity), it is essential that the scientific community works closely with industry, conservation organisations and policy makers to develop robust and efficient conservation and management options.

/pdf/deep-diverse-and-definitely-different-unique-attributes-of-30cfoyr2tg.pdf

Deep, diverse and definitely different: unique attributes of the world's largest ecosystem

The deep sea, the largest ecosystem on Earth and one of the least studied, harbours high biodiversity and provides a wealth of resources. Although humans have used the oceans for millennia, technological developments now allow exploitation of fisheries resources, hydrocarbons and minerals below 2000 m depth. The remoteness of the deep seafloor has promoted the disposal of residues and litter. Ocean acidification and climate change now bring a new dimension of global effects. Thus the challenges facing the deep sea are large and accelerating, providing a new imperative for the science community, industry and national and international organizations to work together to develop successful exploitation management and conservation of the deep-sea ecosystem. This paper provides scientific expert judgement and a semi-quantitative analysis of past, present and future impacts of human-related activities on global deep-sea habitats within three categories: disposal, exploitation and climate change. The analysis is the result of a Census of Marine Life – SYNDEEP workshop (September 2008). A detailed review of known impacts and their effects is provided. The analysis shows how, in recent decades, the most significant anthropogenic activities that affect the deep sea have evolved from mainly disposal (past) to exploitation (present). We predict that from now and into the future, increases in atmospheric CO2 and facets and consequences of climate change will have the most impact on deep-sea habitats and their fauna. Synergies between different anthropogenic pressures and associated effects are discussed, indicating that most synergies are related to increased atmospheric CO2 and climate change effects. We identify deep-sea ecosystems we believe are at higher risk from human impacts in the near future: benthic communities on sedimentary upper slopes, cold-water corals, canyon benthic communities and seamount pelagic and benthic communities. We finalise this review with a short discussion on protection and management methods.

/pdf/man-and-the-last-great-wilderness-human-impact-on-the-deep-2jrgiqly9y.pdf

Man and the last great wilderness: human impact on the deep sea.

Biological structures exert a major influence on species diversity at both local and regional scales on deep continental margins. Some organisms use other species as substrates for attachment, shelter, feeding or parasitism, but there may also be mutual benefits from the association. Here, we highlight the structural attributes and biotic effects of the habitats that corals, sea pens, sponges and xenophyophores offer other organisms. The environmental setting of the biological structures influences their species composition. The importance of benthic species as substrates seems to increase with depth as the complexity of the surrounding geological substrate and food supply decline. There are marked differences in the degree of mutualistic relationships between habitat-forming taxa. This is especially evident for scleractinian corals, which have high numbers of facultative associates (commensals) and few obligate associates (mutualists), and gorgonians, with their few commensals and many obligate associates. Size, flexibility and architectural complexity of the habitat-forming organism are positively related to species diversity for both sessile and mobile species. This is mainly evident for commensal species sharing a facultative relationship with their host. Habitat complexity is enhanced by the architecture of biological structures, as well as by biological interactions. Colony morphology has a great influence on feeding efficiency for suspension feeders. Suspension feeding, habitat-forming organisms modify the environment to optimize their food uptake. This environmental advantage is also passed on to associated filter-feeding species. These effects are poorly understood but represent key points for understanding ecosystems and biodiversity on continental margins. In this paper we explore the contributions of organisms and the biotic structures they create (rather than physical modifications) to habitat heterogeneity and diversity on the deep continental margins.

/pdf/biological-structures-as-a-source-of-habitat-heterogeneity-13wv2fz1nh.pdf

Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins

The Pakistan Margin is characterised by a strong mid-water oxygen minimum zone (OMZ) that intercepts the seabed at bathyal depths (150–1300 m). We investigated whether faunal abundance and diversity trends were similar among protists (foraminiferans and gromiids), metazoan macrofauna and megafauna along a transect (140–1850 m water depth) across the OMZ during the 2003 intermonsoon (March–May) and late/post-monsoon (August–October) seasons. All groups exhibited some drop in abundance in the OMZ core (250–500 m water depth; O2: 0.10–0.13 mL L?1=4.46–5.80 ?M) but to differing degrees. Densities of foraminiferans >63 ?m were slightly depressed at 300 m, peaked at 738 m, and were much lower at deeper stations. Foraminiferans >300 ?m were the overwhelmingly dominant macrofaunal organisms in the OMZ core. Macrofaunal metazoans reached maximum densities at 140 m depth, with additional peaks at 850, 940 and 1850 m where foraminiferans were less abundant. The polychaete Linopherus sp. was responsible for a macrofaunal biomass peak at 950 m. Apart from large swimming animals (fish and natant decapods), metazoan megafauna were absent between 300 and 900 m (O2 0.2 mL L?1=8.92 ?M). The progressively deeper abundance peaks for foraminiferans (>63 ?m), Linopherus sp. and ophiuroids probably represent lower OMZ boundary edge effects and suggest a link between body size and tolerance of hypoxia. Macro- and megafaunal organisms collected between 800 and 1100 m were dominated by a succession of different taxa, indicating that the lower part of the OMZ is also a region of rapid faunal change. Species diversity was depressed in all groups in the OMZ core, but this was much more pronounced for macrofauna and megafauna than for foraminiferans. Oxygen levels strongly influenced the taxonomic composition of all faunal groups. Calcareous foraminiferans dominated the seasonally and permanently hypoxic sites (136–300 m); agglutinated foraminiferans were relatively more abundant at deeper stations where oxygen concentrations were >0.13 mL L?1(=5.80 ?M). Polychaetes were the main macrofaunal taxon within the OMZ; calcareous macrofauna and megafauna (molluscs and echinoderms) were rare or absent where oxygen levels were lowest. The rarity of larger animals between 300 and 700 m on the Pakistan Margin, compared with the abundant macrofauna in the OMZ core off Oman, is the most notable contrast between the two sides of the Arabian Sea. This difference probably reflects the slightly higher oxygen levels and better food quality on the western side.

/pdf/faunal-responses-to-oxygen-gradients-on-the-pakistan-margin-2ydkiexy1i.pdf

Faunal responses to oxygen gradients on the Pakistan margin: A comparison of foraminiferans, macrofauna and megafauna

The genus Gromia includes large marine protists (‘gromiids’) with filose pseudopodia and sack-like organic tests. The first deep-water species were discovered in the 1990s on the Oman Margin of the Arabian Sea and subsequently found on the Pakistan Margin. We present a survey of gromiids in samples collected off Oman in 2002 and off Pakistan in 2003. In addition to the two species (Gromia sphaerica and Gromia pyriformis) already described from this area, at least eight undescribed gromiid species were present. Sausage shaped, grape shaped and spherical morphotypes were represented among this material. On the Oman Margin, gromiids occurred in densities up to several thousand individuals m−2 at 1400 and 1700 m but were much less common at 1100 and 2000 m. Apart from G. pyriformis, which was fairly common (several hundred individuals m−2) at 1000 m, gromiids were uncommon in core samples taken off Pakistan, with 11 indiv. m−2 at 1200 m and 19 indiv. m−2 at 1850 m. On both margins, these protists occurred at depths >1000 m where bottom-water oxygen concentrations exceeded ∼0.2 ml l−1 (=8.92 μM l−1) land sediments were fully bioturbated and oxidised. However, they were not observed at similar oxygen levels above the OMZ. Most gromiids lived on the sediment surface with their apertures facing down and their pseudopodia presumably deployed into the sediment to feed on surficial material and associated bacteria. We conclude that these large protists may play an important ecological role in the bathyal Arabian Sea, particularly in carbon cycling but also in structuring the surficial sediments. In addition, their tests, particularly those of G. sphaerica, provide substrates for attached Foraminifera.

Large organic-walled Protista (Gromia) in the Arabian Sea: Density, diversity, distribution and ecology

Xenophyophores are abundant on a terrace of the lower Nazare Canyon (4300 m water depth) on the Portuguese margin. Here, the most abundant species, Reticulammina cerebreformis sp. nov., occurs in densities of up to 21 individuals/m2. This large species has a soft, friable hemispherical test up to 10 cm in diameter consisting of curved, sinuous plates (lamellae) that branch and anastomose. The plates are separated by deep furrows and other depressions to form a distinctive ‘brain-like’ structure. The outer test layer is thin, weakly cemented and is dominated by fine sediment particles; the internal xenophyae include a higher proportion of larger mineral grains. The second new species at the 4300-m site, Nazareammina tenera gen. et sp. nov., is much less common. The test is basically plate-like, but towards the interior it is perforated by oval spaces, which typically merge into complex system of bar-like features, sometimes with irregular excrescences. The granellare system (cell body and its organic envelope) is packed with tiny mineral grains of various sizes and shapes, including titanium-bearing particles. Also common at this deep site are clusters, with a maximum diameter up to 10 cm or occasionally more, of irregular tubes belonging to Aschemonella ramuliformis Brady 1884, a species previously known mainly from isolated tubes. Rather than being single individuals, these clusters comprise a large number of separate branched tubes. Finally, Syringammina fragillissima Brady 1883, a well-known species that is widely distributed on the NW European margin, occurred on steep sediment-covered slopes at a shallower (1555 m water depth) site in the upper canyon. Almost complete SSU rDNA gene sequences obtained from A. ramuliformis and R. cerebreformis confirm that these previous termxenophyophoresnext term are foraminifera. Together with two previously sequenced xenophophores (Shinkaia lindsayi Lecroq, Gooday, Tsuchiya, Pawlowski 2009 and Syringammina corbicula Richardson 2001), and the foraminiferan Rhizammina algaeformis, they constitute a clade within the radiation of monothalamous foraminifera. In situ photographs of R. cerebreformis and A. ramuliformis reveal no evidence of pseudopodia deployed onto the sediment surface. Instead, these species probably trap sediment within their complex, folded test surface (R. cerebreformis) or intercept suspended particles (A. ramuliformis).

Xenophyophores (Rhizaria, Foraminifera) from the Nazaré Canyon (Portuguese margin, NE Atlantic)

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Faunal responses to oxygen gradients on the Pakistan margin: A comparison of foraminiferans, macrofauna and megafauna

Large organic-walled Protista (Gromia) in the Arabian Sea: Density, diversity, distribution and ecology

Xenophyophores (Rhizaria, Foraminifera) from the Nazaré Canyon (Portuguese margin, NE Atlantic)