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Rabalais, NN, Diaz RJ, Levin LA, Turner RE, Gilbert D, Zhang J.  2010.  Dynamics and distribution of natural and human-caused hypoxia. Biogeosciences. 7:585-619. AbstractWebsite

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 (< 2 mg L(-1), or approximately 30% saturation) can form, therefore, under 'natural' conditions, and are more likely to occur in marine systems when the water residence time is extended, water exchange and ventilation are minimal, stratification occurs, and where carbon production and export to the bottom layer are relatively high. Hypoxia has occurred through geological time and naturally occurs in oxygen minimum zones, deep basins, eastern boundary upwelling systems, and fjords. 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.

Raman, AV, Damodaran R, Levin LA, Ganesh T, Rao YKV, Nanduri S, Madhusoodhanan R.  2015.  Macrobenthos relative to the oxygen minimum zone on the East Indian margin, Bay of Bengal. Marine Ecology-an Evolutionary Perspective. 36:679-700.   10.1111/maec.12176   AbstractWebsite

The Bay of Bengal remains one of the least studied of the world's oxygen minimum zones (OMZs). Here we offer a detailed investigation of the macrobenthos relative to oxygen minimum zone [OMZ - DO (dissolved oxygen), concentration <0.5ml1(-1)] at 110 stations off the North East Indian margin (16(0) and 20(0)N) featuring coastal, shelf and slope settings (10-1004m). Macrobenthos (>0.5mm) composition, abundance and diversity were studied in relation to variations in depth, dissolved oxygen, sediment texture and organic carbon. Using multivariate procedures powered by SIMPROF analysis we identified distinct OMZ core sites (depth 150-280m; DO 0.37ml1(-1)) that exhibited dense populations of surface-feeding polychaetes (mean 2188 ind. m(-2)) represented by spionids and cossurids (96%). Molluscs and crustaceans were poorly represented except for ampeliscid amphipods. The lower OMZ sites (DO>0.55mll(-1)) supported a different assemblage of polychaetes (cirratulids, amphinomids, eunicids, orbinids, paraonids), crustaceans and molluscs, albeit with low population densities (mean 343 ind. m(-2)). Species richness [E(S-100)], diversity (Margalef d; H') and evenness (J') were lower and dominance was higher within the OMZ core region. Multiple regression analysis showed that a combination of sand, clay, organic carbon, and dissolved oxygen explained 62-78% of the observed variance in macrobenthos species richness and diversity: E(S-100) and H'. For polychaetes, clay and oxygen proved important. At low oxygen sites (DO <1mll(-1)), depth accounted for most variance. Residual analysis (after removing depth effects) revealed that dissolved oxygen and sediment organic matter influenced 50-62% of residual variation in E(S-100), H' and d for total macrofauna. Of this, oxygen alone influenced up to similar to 50-62%. When only polychaetes were evaluated, oxygen and organic matter explained up to 58-63%. For low oxygen sites, organic matter alone had the explanatory power when dominance among polychaetes was considered. Overall, macrobenthic patterns in the Bay of Bengal were consistent with those reported for other upwelling margins. However, the compression of faunal gradients at shallower depths was most similar to the Chile/Peru margin, and different from the Arabian Sea, where the depth range of the OMZ is two times greater. The Bay of Bengal patterns may take on added significance as OMZs shoal globally.

Ramirez-Llodra, E, Brandt A, Danovaro R, De Mol B, Escobar E, German CR, Levin LA, Arbizu PM, Menot L, Buhl-Mortensen P, Narayanaswamy BE, Smith CR, Tittensor DP, Tyler PA, Vanreusel A, Vecchione M.  2010.  Deep, diverse and definitely different: unique attributes of the world's largest ecosystem. Biogeosciences. 7:2851-2899.   10.5194/bg-7-2851-2010   AbstractWebsite

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.

Ramirez-Llodra, E, Tyler PA, Baker MC, Bergstad OA, Clark MR, Escobar E, Levin LA, Menot L, Rowden AA, Smith CR, Vandover CL.  2011.  Man and the last great wilderness: human impact on the deep sea. Plos One. 6   10.1371/journal.pone.0022588   AbstractWebsite

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 CO(2) 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 CO(2) 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.

Ramirez-Llodra, E, Trannum HC, Evenset A, Levin LA, Andersson M, Finne TE, Hilario A, Flem B, Christensen G, Schaanning M, Vanreusel A.  2015.  Submarine and deep-sea mine tailing placements: A review of current practices, environmental issues, natural analogs and knowledge gaps in Norway and internationally. Marine Pollution Bulletin. 97:13-35.   10.1016/j.marpolbul.2015.05.062   AbstractWebsite

The mining sector is growing in parallel with societal demands for minerals. One of the most important environmental issues and economic burdens of industrial mining on land is the safe storage of the vast amounts of waste produced. Traditionally, tailings have been stored in land dams, but the lack of land availability, potential risk of dam failure and topography in coastal areas in certain countries results in increasing disposal of tailings into marine systems. This review describes the different submarine tailing disposal methods used in the world in general and in Norway in particular, their impact on the environment (e.g. hyper-sedimentation, toxicity, processes related to changes in grain shape and size, turbidity), current legislation and need for future research. Understanding these impacts on the habitat and biota is essential to assess potential ecosystem changes and to develop best available techniques and robust management plans. (C) 2015 Elsevier Ltd. All rights reserved.

Rasmussen, LL, Cornuelle BD, Levin LA, Largier JL, Di Lorenzo E.  2009.  Effects of small-scale features and local wind forcing on tracer dispersion and estimates of population connectivity in a regional scale circulation model. Journal of Geophysical Research-Oceans. 114   10.1029/2008jc004777   AbstractWebsite

A small-scale model of the Southern California-Northern Baja California coastline has been developed to explore dispersion over the continental shelf, with specific attention to physical parameters pertinent to simulations of larval dispersal and population connectivity. The ROMS simulation employs a nested grid system, with an inner domain resolution of 600 m and an outer domain resolution of 1.5 km. Realistic bathymetry and forcing were employed to investigate the effects of passive transport of tracers introduced at locations with known communities of mytilid mussels along the coastline. The effects of topographic resolution, boundary conditions, and choice of meteorological forcing products on dispersion rates, tracer trajectories, and the subsequent measures of population connectivity were examined. In particular, the choice of wind forcing product resulted in different circulation patterns and tracer trajectories and had especially important consequences on measures of larval connectivity such as self-seeding, potential for larval settlement ( import), and contribution to the pool of available larvae ( export). While some forcing products performed better when model data were compared to field measurements, no product was clearly superior. The uncertainty in results, which may appear minor in larger-scale temperature or surface velocity fields, is significant when examining a sensitive passive tracer. This modeling uncertainty needs to be addressed when interpreting connectivity results.

Rathburn, AE, Levin LA, Tryon M, Gieskes JM, Martin JM, Perez ME, Fodrie FJ, Neira C, Fryer GJ, Mendoza G, McMillan PA, Kluesner J, Adamic J, Ziebis W.  2009.  Geological and biological heterogeneity of the Aleutian margin (1965-4822 m). Progress in Oceanography. 80:22-50.   10.1016/j.pocean.2008.12.002   AbstractWebsite

Geological, biological and biogeochemical characterization of the previously unexplored margin off Unimak Island, Alaska between 1965 and 4822 m water depth was conducted to examine: (1) the geological processes that shaped the margin, (2) the linkages between depth, geomorphology and environmental disturbance in structuring benthic communities of varying size classes and (3) the existence, composition and nutritional sources of methane seep biota on this margin. The study area was mapped and sampled using multibeam sonar, a remotely operated vehicle (ROV) and a towed camera system. Our results provide the first characterization of the Aleutian margin mid and lower slope benthic communities (micro-biota, foraminifera, macrofauna and megafauna), recognizing diverse habitats in a variety of settings. Our investigations also revealed that the geologic feature known as the "Ugamak Slide" is not a slide at all, and could not have resulted from a large 1946 earthquake. However, sediment disturbance appears to be a pervasive feature of this margin. We speculate that the deep-sea occurrence of high densities of Elphidium, typically a shallow-water foraminiferan, results from the influence of sediment redeposition from shallower habitats. Strong representation of cumacean, amphipod and tanaid crustaceans among the Unimak macrofauna may also reflect sediment instability. Although some faunal abundances decline with depth, habitat heterogeneity and disturbance generated by canyons and methane seepage appear to influence abundances of biota in ways that supercede any clear depth gradient in organic matter input. Measures of sediment organic matter and pigment content as well as C and N isotopic signatures were highly heterogeneous, although the availability of organic matter and the abundance of microorganisms in the upper sediment (1-5 cm) were positively correlated. We report the first methane seep on the Aleutian slope in the Unimak region (3263-3285 m), comprised of clam bed, pogonophoran field and carbonate habitats. Seep foraminiferal assemblages were dominated by agglutinated taxa, except for habitats above the seafloor on pogonophoran tubes. Numerous infaunal taxa in clam bed and pogonophoran field sediments and deep-sea "reef' cnidarians (e.g., corals and hydroids) residing on rocks near seepage sites exhibited light organic delta(13)C signatures indicative of chemosynthetic nutritional sources. The extensive geological, biogeochemical and biological heterogeneity as well as disturbance features observed on the Aleutian slope provide an attractive explanation for the exceptionally high biodiversity characteristic of the world's continental margins. (C) 2008 Elsevier Ltd. All rights reserved.

Rathburn, AE, Levin LA, Held Z, Lohmann KC.  2000.  Benthic foraminifera associated with cold methane seeps on the northern California margin: Ecology and stable isotopic composition. Marine Micropaleontology. 38:247-266.   10.1016/s0377-8398(00)00005-0   AbstractWebsite

Release of methane from large marine reservoirs has been linked to climate change, as a causal mechanism and a consequence of temperature changes, during the Quaternary and the Paleocene. These inferred linkages are based primarily on variations in benthic foraminiferal stable isotope signatures. Few modem analog data exist, however, to assess the influence of methane flux on the geochemistry or faunal characteristics of benthic foraminiferal assemblages. Here we present analyses of the ecology and stable isotopic compositions of living (Rose Bengal stained) and dead (fossil) foraminifera (>150 mu m) from cold methane seeps on the slope off of the Eel River, northern California (500-525 m), and discuss potential applications for reconstructions of methane release in the past and present. Calcareous foraminiferal assemblages associated with Calyptogena clam bed seeps were comprised of species that are also found in organic-rich environments. Cosmopolitan, paleoceanographically important taxa were abundant; these included Uvigerina, Bolivina, Chilostomella, Globobulimina, and Nonionella. We speculate that seep foraminifera are attracted to the availability of food at cold seeps, and require no adaptations beyond those needed for life in organic-rich, reducing environments. Oxygen isotopic values of the tests of living foraminiferal assemblages from seeps had a high range (up to 0.69 parts per thousand) as did carbon isotopic values (up to 1.02 parts per thousand). Many living foraminiferal isotope values were within the range exhibited by the same or similar species in non-seep environments. Carbon isotopic values of fossil foraminifera found deeper in the sediments (18-20 cm), however, were 4.10 parts per thousand (U. peregrina) and 3.60 parts per thousand (B. subargentea) more negative than living delta(13)C values. These results suggest that delta(13)C values of foraminiferal tests reflect methane seepage and species-specific differences in isotopic composition, and can indicate temporal variations in seep activity. A better understanding of foraminiferal ecology and stable isotopic composition will enhance paleo-seep recognition, and improve interpretations of climatic and paleoceanographic change. (C) 2000 Elsevier Science B.V. All rights reserved.

Rice, SA, Levin LA.  1998.  Streblospio gynobranchiata, a new spionid polychaete species (Annelida : Polychaeta) from Florida and the Gulf of Mexico with an analysis of phylogenetic relationships within the genus Streblospio. Proceedings of the Biological Society of Washington. 111:694-707. AbstractWebsite

Streblospio gynobranchiata, a new species of spionid polychaete, is described from Florida and the Gulf of Mexico. This species is distinguished by the presence of strap-like branchiae, beneath which larvae are brooded, in the genital body region of the females. Data from experimental crosses conducted in the laboratory are presented that indicate incipient reproductive isolation between S. gynobranchiata from Florida and S. benedicti from North Carolina and California. The spermatophores produced by males of S. gynobranchiata and S. benedicti are described for the first time. Morphological, developmental, and reproductive characteristics are used in combination to construct a cladogram that suggests possible phylogenetic relationships between known species in the genus Streblospio.

Robinson, CA, Bernhard JM, Levin LA, Mendoza GF, Blanks JK.  2004.  Surficial hydrocarbon seep infauna from the Blake Ridge (Atlantic Ocean, 2150 m) and the Gulf of Mexico (690-2240 m). Marine Ecology-Pubblicazioni Della Stazione Zoologica Di Napoli I. 25:313-336.   10.1111/j.1439-0485.2004.00034.x   AbstractWebsite

Infauna, including foraminifera and metazoans, were enumerated and identified from five types of seep habitats and two adjacent non-seep habitats. Collections were made with the deep submergence research vessel 'Alvin' from three areas of active seepage in the Gulf of Mexico (Alaminos Canyon [2220 m], Atwater Canyon [1930 m], and Green Canyon lease block 272 [700 m]) and on the Blake Ridge Diapir [2250 m], which is located off the southeastern coast of the United States. The seep habitats sampled included four types of microbial mats (Beggiatoa, Thioploca, thin and thick Arcobacter) and the periphery of a large mussel bed. Sediments under large rhizopod protists, xenophyophores, were sampled adjacent to the mussel bed periphery. A non-seep site, which was >1 km away from active seeps, was also sampled for comparison. Densities of most taxa were higher in the Gulf of Mexico seeps than in Blake Ridge samples, largely because densities in the thick microbial mats of Blake Ridge were significantly lower. Diversity was higher in the Thioploca mats compared to other microbial-mat types. Within an ocean basin (i.e., Atlantic, Gulf of Mexico) we did not observe significant differences in meiofaunal or macrofaunal composition in Beggiatoa versus Thioploca mats or thin versus thick Arcobacter mats. Foraminifera represented up to 16% of the seep community, a proportion that is comparable to their contribution at adjacent non-seep communities. In general, the observed densities and taxonomic composition of seep sites at the genus level was consistent with previous observations from seeps (e.g., the foraminifers Bolivina and Fursenkoina, the dorvilleid polychaete Ophryotrocha).