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2019
Levin, LA, Bett BJ, Gates AR, Heimbach P, Howe BM, Janssen F, McCurdy A, Ruhl HA, Snelgrove P, Stocks KI, Bailey D, Baumann-Pickering S, Beaverson C, Benfield MC, Booth DJ, Carreiro-Silva M, Colaco A, Eble MC, Fowler AM, Gjerde KM, Jones DOB, Katsumata K, Kelley D, Le Bris N, Leonardi AP, Lejzerowicz F, Macreadie PI, McLean D, Meitz F, Morato T, Netburn A, Pawlowski J, Smith CR, Sun S, Uchida H, Vardaro MF, Venkatesan R, Weller RA.  2019.  Global observing needs in the deep ocean. Frontiers in Marine Science. 6   10.3389/fmars.2019.00241   AbstractWebsite

The deep ocean below 200 m water depth is the least observed, but largest habitat on our planet by volume and area. Over 150 years of exploration has revealed that this dynamic system provides critical climate regulation, houses a wealth of energy, mineral, and biological resources, and represents a vast repository of biological diversity. A long history of deep-ocean exploration and observation led to the initial concept for the Deep-Ocean Observing Strategy (DOOS), under the auspices of the Global Ocean Observing System (GOOS). Here we discuss the scientific need for globally integrated deep-ocean observing, its status, and the key scientific questions and societal mandates driving observing requirements over the next decade. We consider the Essential Ocean Variables (EOVs) needed to address deep-ocean challenges within the physical, biogeochemical, and biological/ecosystem sciences according to the Framework for Ocean Observing (FOO), and map these onto scientific questions. Opportunities for new and expanded synergies among deep-ocean stakeholders are discussed, including academic-industry partnerships with the oil and gas, mining, cable and fishing industries, the ocean exploration and mapping community, and biodiversity conservation initiatives. Future deep-ocean observing will benefit from the greater integration across traditional disciplines and sectors, achieved through demonstration projects and facilitated reuse and repurposing of existing deep-sea data efforts. We highlight examples of existing and emerging deep-sea methods and technologies, noting key challenges associated with data volume, preservation, standardization, and accessibility. Emerging technologies relevant to deep-ocean sustainability and the blue economy include novel genomics approaches, imaging technologies, and ultra-deep hydrographic measurements. Capacity building will be necessary to integrate capabilities into programs and projects at a global scale. Progress can be facilitated by Open Science and Findable, Accessible, Interoperable, Reusable (FAIR) data principles and converge on agreed to data standards, practices, vocabularies, and registries. We envision expansion of the deep-ocean observing community to embrace the participation of academia, industry, NGOs, national governments, international governmental organizations, and the public at large in order to unlock critical knowledge contained in the deep ocean over coming decades, and to realize the mutual benefits of thoughtful deep-ocean observing for all elements of a sustainable ocean.

2012
Levin, LA, Orphan VJ, Rouse GW, Rathburn AE, Ussler W, Cook GS, Goffredi SK, Perez EM, Waren A, Grupe BM, Chadwick G, Strickrott B.  2012.  A hydrothermal seep on the Costa Rica margin: middle ground in a continuum of reducing ecosystems. Proceedings of the Royal Society B-Biological Sciences. 279:2580-2588.   10.1098/rspb.2012.0205   AbstractWebsite

Upon their initial discovery, hydrothermal vents and methane seeps were considered to be related but distinct ecosystems, with different distributions, geomorphology, temperatures, geochemical properties and mostly different species. However, subsequently discovered vents and seep systems have blurred this distinction. Here, we report on a composite, hydrothermal seep ecosystem at a subducting seamount on the convergent Costa Rica margin that represents an intermediate between vent and seep ecosystems. Diffuse flow of shimmering, warm fluids with high methane concentrations supports a mixture of microbes, animal species, assemblages and trophic pathways with vent and seep affinities. Their coexistence reinforces the continuity of reducing environments and exemplifies a setting conducive to interactive evolution of vent and seep biota.

Levin, LA, Sibuet M.  2012.  Understanding Continental Margin Biodiversity: A New Imperative. Annual Review of Marine Science, Vol 4. 4( Carlson CA, Giovannoni SJ, Eds.).:79-+., Palo Alto: Annual Reviews   10.1146/annurev-marine-120709-142714   Abstract

Until recently, the deep continental margins (200-4,000 m) were perceived as monotonous mud slopes of limited ecological or environmental concern. Progress in seafloor mapping and direct observation now reveals unexpected heterogeneity, with a mosaic of habitats and ecosystems linked to geomorphological, geochemical, and hydrographic features that influence biotic diversity. Interactions among water masses, terrestrial inputs, sediment diagenesis, and tectonic activity create a multitude of ecological settings supporting distinct communities that populate canyons and seamounts, high-stress oxygen minimum zones, and methane seeps, as well as vast reefs of cold corals and sponges. This high regional biodiversity is fundamental to the production of valuable fisheries, energy, and mineral resources, and performs critical ecological services (nutrient cycling, carbon sequestration, nursery and habitat support). It is under significant threat from climate change and human resource extraction activities. Serious actions are required to preserve the functions and services provided by the deep-sea settings we are just now getting to know.

2010
Buhl-Mortensen, L, Vanreusel A, Gooday AJ, Levin LA, Priede IG, Buhl-Mortensen P, Gheerardyn H, King NJ, Raes M.  2010.  Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. Marine Ecology-an Evolutionary Perspective. 31:21-50.   10.1111/j.1439-0485.2010.00359.x   AbstractWebsite

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.

2009
Basak, C, Rathburn AE, Perez ME, Martin JB, Kluesner JW, Levin LA, De Deckker P, Gieskes JM, Abriani M.  2009.  Carbon and oxygen isotope geochemistry of live (stained) benthic foraminifera from the Aleutian Margin and the Southern Australian Margin. Marine Micropaleontology. 70:89-101.   10.1016/j.marmicro.2008.11.002   AbstractWebsite

Comparisons of ambient bottom-water geochemistry and stable isotopic values of the tests of living (stained) calcareous benthic foraminifera from the North Pacific (on the Aleutian Margin, water depth 1988 m) and Murray Canyons group in the Southern Indian Ocean (Australian Margin, water depths 2476 m and 1634 m) provide modem environmental analogs to calibrate paleoenvironmental assessments. Consistent with the hypothesis that microhabitat preferences influence foraminiferal isotopic values, benthic foraminifera from both margins were depleted in (13)C with respect to bottom-water dissolved inorganic carbon (DIC). The carbon isotope values of deep infaunal foraminifera (Chilostomella oolina, Globobulimina pacifica) showed greater differences from estimates of those of DIC than shallow benthic foraminifera (Bulimina mexicana, Bolivinita quadrilatera, Pullenia bulloides). This study provides new isotopic and ecological information for B. quadrilatera. The mean Delta delta(13)C value, defined as foraminiferal delta(13)C values minus estimated ambient delta(13)C values from the Aleutian Margin, is 0.97 parts per thousand higher for G. pacifica than the mean from the Murray Canyon. This difference may result either from genetic or biological differences between the populations or from differences in environmental isotopic influences (such as pore water differences) that were not accounted for in the equilibrium calculations. These analyses provide calibration information for the evaluation of bottom water conditions and circulation patterns of ancient oceans based on fossil foraminiferal geochemistry. (C) 2008 Elsevier B.V. All rights reserved.

Woulds, C, Andersson JH, Cowie GL, Middelburg JJ, Levin LA.  2009.  The short-term fate of organic carbon in marine sediments: Comparing the Pakistan margin to other regions. Deep Sea Research (Part II, Topical Studies in Oceanography). 56:393-402., United Kingdom: Elsevier BV   10.1016/j.dsr2.2008.10.008   AbstractWebsite

Pulse-chase experiments with isotopically labelled phytodetritus conducted across the Pakistan margin reveal that the impact of biological activities on benthic C-cycling varies markedly among sites exhibiting different seafloor conditions. In this study, patterns of biological C-processing across the Pakistan margin oxygen minimum zone (OMZ) are compared with those observed in previous tracer studies. Variations in site environmental conditions are proposed to explain the considerable variations in C-processing patterns among this and previous studies. Three categories of C-processing pattern are identified: (1) respiration dominated, where respiration accounts for >75% of biological C-processing, and uptake by metazoan macrofauna, foraminifera and bacteria are relatively minor processes. These sites tend to show several (although not necessarily all) of the properties of being cold and deep, and having low inputs of organic carbon to the sediment and relatively low-biomass metazoan macrofaunal communities; (2) active faunal uptake, where respiration accounts for <75%, and metazoan macrofaunal, foraminiferal and bacterial uptake each account for 10-25% of biological C-processing. This type is further split into metazoan macrofaunal- and foraminiferal-dominated situations, dictated by oxygen availability; and (3) metazoan macrofaunal uptake dominated, characterised by metazoan macrofaunal uptake accounting for ~50% of biological C-processing, due to unusually large biomasses of the phytodetritus-consuming animals. Total respiration rates (of added C) on the Pakistan margin fell within the range of rates measured elsewhere in the deep sea (} .1-2.8mgCm super(-) super(2)h super(-) super(1)), and seem to be dominantly controlled by seafloor temperature. Rates of metazoan macrofaunal uptake of organic matter (OM) on the Pakistan margin are larger than those in most other studies, and this is attributed to the large and active metazoan macrofaunal communities in the lower OMZ, characteristic of OMZ boundaries. Finally, biological mixing of Pakistan margin sediments was reduced compared to that observed in comparable tracer studies on other margins. This probably reflects faunal feeding and burrowing strategies consistent with low oxygen concentrations and a relatively abundant supply of sedimentary OM.

2006
Arntz, WE, Gallardo VA, Gutierrez D, Isla E, Levin LA, Mendo J, Neira C, Rowe GT, Tarazona J, Wolff M.  2006.  El NiƱo and similar perturbation effects on the benthos of the Humboldt, California, and Benguela Current upwelling ecosystems. Advances in Geosciences. 6:243-265.: European Geosciences Union, c/o E.O.S.T. 5, rue Rene Descartes Strasbourg Cedex 67084 France, [mailto:egu.production@copernicus.org], [URL:http://www.copernicus.org/EGU] AbstractWebsite

To a certain degree, Eastern Boundary Current (EBC) ecosystems are similar: Cold bottom water from moderate depths, rich in nutrients, is transported to the euphotic zone by a combination of trade winds, Coriolis force and Ekman transport. The resultant high primary production fuels a rich secondary production in the upper pelagic and nearshore zones, but where O sub(2) exchange is restricted, it creates oxygen minimum zones (OMZs) at shelf and upper slope (Humboldt and Benguela Current) or slope depths (California Current). These hypoxic zones host a specifically adapted, small macro- and meiofauna together with giant sulphur bacteria that use nitrate to oxydise H sub(2)S. In all EBC, small polychaetes, large nematodes and other opportunistic benthic species have adapted to the hypoxic conditions and co-exist with sulphur bacteria, which seem to be particularly dominant off Peru and Chile. However, a massive reduction of macrobenthos occurs in the core of the OMZ. In the Humboldt Current area the OMZ ranges between <100 and about 600 m, with decreasing thickness in a poleward direction. The OMZ merges into better oxygenated zones towards the deep sea, where large cold-water mega- and macrofauna occupy a dominant role as in the nearshore strip. The Benguela Current OMZ has a similar upper limit but remains shallower. It also hosts giant sulphur bacteria but little is known about the benthic fauna. However, sulphur eruptions and intense hypoxia might preclude the coexistence of significant mega- und macrobenthos. Conversely, off North America the upper limit of the OMZ is considerably deeper (e.g., 500-600 m off California and Oregon), and the lower boundary may exceed 1000m. The properties described are valid for very cold and cold (La Nina and "normal") ENSO conditions with effective upwelling of nutrient-rich bottom water. During warm (El Nino) episodes, warm water masses of low oxygen concentration from oceanic and equatorial regions enter the upwelling zones, bringing a variety of (sub)tropical immigrants. The autochthonous benthic fauna emigrates to deeper water or poleward, or suffers mortality. However, some local macrofaunal species experience important population proliferations, presumably due to improved oxygenation (in the southern hemisphere), higher temperature tolerance, reduced competition or the capability to use different food. Both these negative and positive effects of el Nino influence local artisanal fisheries and the livelihood of coastal populations. In the Humboldt Current system the hypoxic seafloor at outer shelf depths receives important flushing from the equatorial zone, causing havoc on the sulphur bacteria mats and immediate recolonisation of the sediments by mega- and macrofauna. Conversely, off California, the intruding equatorial water masses appear to have lower oxygen than ambient waters, and may cause oxygen deficiency at upper slope depths. Effects of this change have not been studied in detail, although shrimp and other taxa appear to alter their distribution on the continental margin. Other properties and reactions of the two Pacific EBC benthic ecosystems to el Nino seem to differ, too, as does the overall impact of major episodes (e.g., 1982/1983(1984) vs. 1997/1998). The relation of the "Benguela Nino" to ENSO seems unclear although many Pacific- Atlantic ocean and atmosphere teleconnections have been described. Warm, low- oxygen equatorial water seems to be transported into the upwelling area by similar mechanisms as in the Pacific, but most major impacts on the eukaryotic biota obviously come from other, independent perturbations such as an extreme eutrophication of the sediments ensuing in sulphidic eruptions and toxic algal blooms. Similarities and differences of the Humboldt and California Current benthic ecosystems are discussed with particular reference to ENSO impacts since 1972/73. Where there are data available, the authors include the Benguela Current ecosystem as another important, non-Pacific EBC, which also suffers from the effects of hypoxia.

1999
Thistle, D, Levin LA, Gooday AJ, Pfannkuche O, Lambshead PJD.  1999.  Physical reworking by near-bottom flow alters the metazoan meiofauna of Fieberling Guyot (northeast Pacific). Deep-Sea Research Part I-Oceanographic Research Papers. 46:2041-2052.   10.1016/s0967-0637(99)00040-0   AbstractWebsite

Although much of the deep sea is physically tranquil, some regions experience near-bottom flows that rework the surficial sediment. During periods of physical reworking, animals in the reworked layer risk being suspended, which can have both positive and negative effects. Reworking can also change the sediment in ecologically important ways, so the fauna of reworked sites should differ from that of quiescent locations. We combined data from two reworked, bathyal sites on the summit of Fieberling Guyot (32 degrees 27.631'N, 127 degrees 49.489'W; 32 degrees 27.581'N, 127 degrees 47.839'W) and compared the results with those of more tranquil sites. We tested for differences in the following parameters, which seemed likely to be sensitive to the direct or indirect effects of reworking: (1) the vertical distribution of the meiofauna in the sea bed, (2) the relative abundance of surface-living harpacticoids, (3) the proportion of the fauna consisting of interstitial harpacticoids, (4) the ratio of harpacticoids to nematodes. We found that the vertical distributions of harpacticoid copepods, ostracods, and kinorhynchs were deeper on Fieberling. In addition, the relative abundance of surface-living harpacticoids was less, the proportion of interstitial harpacticoids was greater, and the ratio of harpacticoids to nematodes was greater on Fieberling. These differences between Fieberling and the comparison sites suggest that physical reworking affects deep-sea meiofauna and indicate the nature of some of the effects. (C) 1999 Elsevier Science Ltd. AII rights reserved.

1998
Thistle, D, Levin LA.  1998.  The effect of experimentally increased near-bottom flow on metazoan meiofauna at a deep-sea site, with comparison data on macrofauna. Deep-Sea Research Part I-Oceanographic Research Papers. 45:625-+.   10.1016/s0967-0637(97)00101-5   AbstractWebsite

It has been argued that strong near-bottom hows affect macrofauna and meiofauna in the deep sea, but the evidence comes largely from studies that compared sites separated geographically by hundreds to thousands of kilometers and in depth by hundreds of meters. In this paper, the results of the first experimental investigation of the effects of strong near-bottom flow on deep-sea metazoan meiofauna are presented. At a site (32 degrees 27.581' N, 127 degrees 47.839' W) at 583 m depth on the Fieberling Guyot summit plain, the submersible Alvin emplaced weirs designed to increase the near-bottom flow locally. After 6.5 weeks, sediments in the weirs and unmanipulated locations in the vicinity were sampled. The abundances of nematodes, harpacticoid copepods, ostracods, and kinorhynchs, considered collectively and as individual taxa, were significantly lower in the weir samples than in the background samples. Parallel responses were observed in total macrofaunal and mollusk abundances. Proportional declines in kinorhynchs and mollusks were observed as well. These results suggest that strong near-bottom flow can reduce the abundance of meiofauna and macrofauna in the deep sea and alter assemblage composition. (C) 1998 Elsevier Science Ltd. All rights reserved.

1991
Levin, LA, McCann LD, Thomas CL.  1991.  The ecology of polychaetes on deep seamounts in the eastern Pacific Ocean. Ophelia. :467-476. AbstractWebsite

Polychaetes were collected by the submersible ALVIN on 18 deep (788-3,353 m) seamounts in the eastern Pacific Ocean at 10-degrees, 13-degrees, 20-degrees and 30-degrees N off western Mexico. Polychaetes comprised 57.7% of all macrofauna collected. Average density over all locations was 942 polychaetes/m2. Thirty-eight families were represented among the 1,422 infaunal polychaetes collected. Five families, the Paraonidae, Cirratulidae, Syllidae, Ampharetidae, and Sabellidae, attained average densities > 1 individual/196 cm2 core. We evaluated effects of latitude, local setting, depth, and substrate on polychaete abundance, taxonomic composition, and lifestyles. Unusually high polychaete densities (7,194/m2) and low diversities were observed in a shallow caldera (788 m) at 13-degrees N. Excluding this site, the latitude exhibiting the highest polychaete densities (xBAR = 939/m2) was 10-degrees N. Of the seven settings examined, pit craters (within seamount calderas) supported the highest densities (xBAR = 1031/m2), and hydrothermal oxide fields and seamount bases exhibited the lowest polychaete densities (xBAR = 576-612/m2). Rippled foraminiferal sands on volcano summits supported large numbers of filter feeders, particularly sabellids. Regressions of total polychaete abundance on depth and on percent sand were not significant. Large, epifaunal, sediment-agglutinating protozoans (Phylum Sarcodina: Class Xenophyophorea) provided habitat for 34 polychaete species. Polychaete abundance and family composition were generally similar to those reported for other nearshore, deep-sea environments at comparable depths. With the exception of the shallowest site, species richness was typically high.