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2016
Burkett, AM, Rathburn AE, Perez ME, Levin LA, Martin JB.  2016.  Colonization of over a thousand Cibicidoides wuellerstorfi (foraminifera: Schwager, 1866) on artificial substrates in seep and adjacent off-seep locations in dysoxic, deep-sea environments. Deep-Sea Research Part I-Oceanographic Research Papers. 117:39-50.   10.1016/j.dsr.2016.08.011   AbstractWebsite

After-1 yr on the seafloor at water depths of similar to 700 m on Hydrate Ridge in the Pacific, eight colonization experiments composed primarily of a plastic mesh cube (from here on refered to as SEA(3), for Seafloor Epibenthic Attachment Cubes) were colonized by 1076 Cibicidoides wuellerstorfi on similar to 1841 cm(2) of experimental substrate. This species is typically considered an indicator of well-oxygenated conditions, and recruitment of such large numbers in bottom waters with low dissolved oxygen availability (0.24-0.37 mL/L) indicate that this taxon may not be as limited by oxygen as previously thought. Clues about substrate preferences were evident from the distribution, or lack thereof, of individuals among plastic mesh, coated steel frame, wooden dowels and reflective tape. Abundance, individual size distributions within cage populations and isotopic biogeochemistry of living foraminifera colonizing experimental substrates were compared between active seep and adjacent off seep experiment locations, revealing potential differences between these environments. Few studies have examined foraminiferal colonization of hard substrates in the deep-sea and to our knowledge no previous study has compared foraminiferal colonization of active seep and off-seep substrates from the same region. This study provides initial results of recruitment, colonization, geochemical and morphological aspects of the paleoceanographically significant species, C. wuellerstorfi, from dynamic deep-sea environments. Further experimental deployments of SEA(3)s will provide a means to assess relatively unknown ecologic dynamics of important foraminiferal deep-sea species.

2015
Burkett, AM, Rathburn AE, Perez ME, Levin LA, Cha H, Rouse GW.  2015.  Phylogenetic placement of Cibicidoides wuellerstorfi (Schwager, 1866) from methane seeps and non-seep habitats on the Pacific margin. Geobiology. 13:44-52.   10.1111/gbi.12118   AbstractWebsite

Benthic foraminifera are among the most abundant groups found in deep-sea habitats, including methane seep environments. Unlike many groups, no endemic foraminiferal species have been reported from methane seeps, and to our knowledge, genetic data are currently sparse for Pacific deep-sea foraminifera. In an effort to understand the relationships between seep and non-seep populations of the deep-sea foraminifera Cibicidoides wuellerstorfi, a common paleo-indicator species, specimens from methane seeps in the Pacific were analyzed and compared to one another for genetic similarities of small subunit rDNA (SSU rDNA) sequences. Pacific Ocean C. wuellerstorfi were also compared to those collected from other localities around the world (based on 18S gene available on Genbank, e.g., Schweizer et al., 2009). Results from this study revealed that C. wuellerstorfi living in seeps near Costa Rica and Hydrate Ridge are genetically similar to one another at the species level. Individuals collected from the same location that display opposite coiling directions (dextral and sinstral) had no species level genetic differences. Comparisons of specimens with genetic information available from Genbank (SSU rDNA) showed that Pacific individuals, collected for this study, are genetically similar to those previously analyzed from the North Atlantic and Antarctic. These observations provide strong evidence for the true cosmopolitan nature of C. wuellerstorfi and highlight the importance of understanding how these microscopic organisms are able to maintain sufficient genetic exchange to remain within the same species between seep and non-seep habitats and over global distances.

2014
Larkin, KE, Gooday AJ, Woulds C, Jeffreys RM, Schwartz M, Cowie G, Whitcraft C, Levin L, Dick JR, Pond DW.  2014.  Uptake of algal carbon and the likely synthesis of an "essential" fatty acid by Uvigerina ex. gr. semiornata (Foraminifera) within the Pakistan margin oxygen minimum zone: evidence from fatty acid biomarker and C-13 tracer experiments. Biogeosciences. 11:3729-3738.   10.5194/bg-11-3729-2014   AbstractWebsite

Foraminifera are an important component of benthic communities in oxygen-depleted settings, where they potentially play a significant role in the processing of organic matter. We tracked the uptake of a C-13-labelled algal food source into individual fatty acids in the benthic foraminiferal species Uvigerina ex. gr. semiornata from the Arabian Sea oxygen minimum zone (OMZ). The tracer experiments were conducted on the Pakistan margin during the late/post monsoon period (August-October 2003). A monoculture of the diatom Thalassiosira weisflogii was C-13-labelled and used to simulate a pulse of phytoplankton in two complementary experiments. A lander system was used for in situ incubations at 140m water depth and for 2.5 days in duration. Shipboard laboratory incubations of cores collected at 140 m incorporated an oxystat system to maintain ambient dissolved oxygen concentrations and were terminated after 5 days. Uptake of diatoms was rapid, with a high incorporation of diatom fatty acids into foraminifera after similar to 2 days in both experiments. Ingestion of the diatom food source was indicated by the increase over time in the quantity of diatom biomarker fatty acids in the foraminifera and by the high percentage of C-13 in many of the fatty acids present at the endpoint of both in situ and laboratory-based experiments. These results indicate that U. ex. gr. semiornata rapidly ingested the diatom food source and that these foraminifera will play an important role in the short-term cycling of organic matter within this OMZ environment. The presence of 18:1(n-7) in the experimental foraminifera suggested that U. ex. gr. semiornata also consumed non-labelled bacterial food items. In addition, levels of 20:4(n-6), a PUFA only present in low amounts in the diatom food, increased dramatically in the foraminifera during both the in situ and shipboard experiments, possibly because it was synthesised de novo. This "essential fatty acid" is often abundant in benthic fauna, yet its origins and function have remained unclear. If U. ex. gr. semiornata is capable of de novo synthesis of 20:4(n-6), then it represents a potentially major source of this dietary nutrient in benthic food webs.

2013
Levin, LA, McGregor AL, Mendoza GF, Woulds C, Cross P, Witte U, Gooday AJ, Cowie G, Kitazato H.  2013.  Macrofaunal colonization across the Indian margin oxygen minimum zone. Biogeosciences. 10:7161-7177.   10.5194/bg-10-7161-2013   AbstractWebsite

There is a growing need to understand the ability of bathyal assemblages to recover from disturbance and oxygen stress, as human activities and expanding oxygen minimum zones increasingly affect deep continental margins. The effects of a pronounced oxygen minimum zone (OMZ) on slope benthic community structure have been studied on every major upwelling margin; however, little is known about the dynamics or resilience of these benthic populations. To examine the influence of oxygen and phytodetritus on shortterm settlement patterns, we conducted colonization experiments at 3 depths on the West Indian continental margin. Four colonization trays were deployed at each depth for 4 days at 542 and 802 m (transect 1-16 degrees 58 ' N) and for 9 days at 817 and 1147 m (transect 2-17 degrees 31 ' N). Oxygen concentrations ranged from 0.9 mu M (0.02 mLL(-1)) at 542 m to 22 mu M (0.5 mLL(-1) ) at 1147 m. All trays contained local defaunated sediments; half of the trays at each depth also contained C-13/N-15-labeled phytodetritus mixed into the sediments. Sediment cores were collected between 535 m and 1140 m from 2 cross-margin transects for analysis of ambient (source) macrofaunal (> 300 mu m) densities and composition. Ambient macrofaunal densities ranged from 0 ind m(-2) (at 535-542 m) to 7400 ind m(-2), with maximum values on both transects at 700-800 m. Macrofaunal colonizer densities ranged from 0 ind m(-2) at 542 m, where oxygen was lowest, to average values of 142 ind m(-2) at 800 m, and 3074 ind m(-2) at 1147 m, where oxygen concentration was highest. These were equal to 4.3 and 151% of the ambient community at 800 m and 1147 m, respectively. Community structure of settlers showed no response to the presence of phytodetritus. Increasing depth and oxygen concentration, however, significantly influenced the community composition and abundance of colonizing macrofauna. Polychaetes constituted 92.4% of the total colonizers, followed by crustaceans (4.2%), mollusks (2.5%), and echinoderms (0.8%). The majority of colonizers were found at 1147 m; 88.5% of these were Capitella sp., although they were rare in the ambient community. Colonists at 800 and 1147 m also included ampharetid, spionid, syllid, lumbrinerid, cirratulid, cossurid and sabellid polychaetes. Consumption of C-13/N-15-labeled phytodetritus was observed for macrofaunal foraminifera (too large to be colonizers) at the 542 and 802/817 m sites, and by metazoan macrofauna mainly at the deepest, better oxygenated sites. Calcareous foraminifera (Uvigerina, Hoeglundina sp.), capitellid polychaetes and cumaceans were among the major phytodetritus consumers. These preliminary experiments suggest that bottom-water oxygen concentrations may strongly influence ecosystem services on continental margins, as reflected in rates of colonization by benthos and colonizer processing of carbon following disturbance. They may also provide a window into future patterns of settlement on the continental slope as the world's oxygen minimum zones expand.

2012
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.

2011
Bailey, JV, Salman V, Rouse GW, Schulz-Vogt HN, Levin LA, Orphan VJ.  2011.  Dimorphism in methane seep-dwelling ecotypes of the largest known bacteria. ISME Journal. 5:1926-1935.   10.1038/ismej.2011.66   AbstractWebsite

We present evidence for a dimorphic life cycle in the vacuolate sulfide-oxidizing bacteria that appears to involve the attachment of a spherical Thiomargarita-like cell to the exteriors of invertebrate integuments and other benthic substrates at methane seeps. The attached cell elongates to produce a stalk-like form before budding off spherical daughter cells resembling free-living Thiomargarita that are abundant in surrounding sulfidic seep sediments. The relationship between the attached parent cell and free-living daughter cell is reminiscent of the dimorphic life modes of the prosthecate Alphaproteobacteria, but on a grand scale, with individual elongate cells reaching nearly a millimeter in length. Abundant growth of attached Thiomargarita-like bacteria on the integuments of gastropods and other seep fauna provides not only a novel ecological niche for these giant bacteria, but also for animals that may benefit from epibiont colonization. The ISME Journal (2011) 5, 1926-1935; doi: 10.1038/ismej.2011.66; published online 23 June 2011

2010
Levin, LA, Mendoza GF, Gonzalez JP, Thurber AR, Cordes EE.  2010.  Diversity of bathyal macrofauna on the northeastern Pacific margin: the influence of methane seeps and oxygen minimum zones. Marine Ecology-an Evolutionary Perspective. 31:94-110.   10.1111/j.1439-0485.2009.00335.x   AbstractWebsite

The upper continental slope in the northeastern Pacific Ocean is intercepted by a deep oxygen minimum zone (OMZ; 650-1100 m) and punctuated by conduits of methane seepage. We examined the effects of these two dominant sources of heterogeneity on the density, composition and diversity of heterotrophic macrofauna off Hydrate Ridge, Oregon (OR; 800 m water depth), where the seeps co-occur within an OMZ, and off the Eel River, Northern California (CA; 500 m), where seeps are overlain by better oxygenated waters. We hypothesized that seeps (containing clam beds and microbial mats) should contribute a suite of distinct species to the regional margin species pool but that OMZ-associated hypoxia would dampen seep-related heterogeneity. Macrofaunal densities were highest (23,000-33,510 ind.m(-2)) in the CA seep sediments and in the OR near-seep samples, intermediate in the OR seep, CA near seep and CA and OR 500-m margin sediments (10,05419,777 ind.m(-2)), and lowest in the CA and OR OMZ habitats at 800 m (42697847 ind.m(-2)). Annelids constituted over 50% of the taxa in all but the CA clam bed and OR microbial mat sediments, where mollusks were abundant. Approximately 50% of seep species appeared to be habitat endemic; species present in microbial mats largely formed a subset of those present in the clam beds. Dorvilleid and ampharetid polychaetes were dominant in the seep sediments; non-seep margin sediments at 500 and 800 m were populated heavily by branckiate polychaetes including cossurids and paraonids. Alpha diversity (Es[20] calculated per core) was lowest and rank 1 dominance was highest in the CA and OR microbial mat habitats. Pooled analyses of Es[100] revealed highest species richness in the CA clam bed and near-seep habitats (30.3 and 29.6, respectively), and lowest species richness in the OR microbial mat and near-seep habitats (16.5 and 17.9, respectively). Non-seep sediments (500 and 800 m) off both CA and OR were more homogeneous (55-57% within-habitat similarity) than clam bed and microbial mat sediments (only 32-37% within-habitat similarity). CA sediment macrofauna generally exhibit higher alpha diversity, and as habitats are combined, a higher rate of increase in the slope of the species accumulation curves than do OR margin macrofauna. Methane seeps in the NE Pacific introduce significant heterogeneity that increases margin biodiversity at multiple spatial scales. However, our hypothesis that the OMZ would lessen the seep contributions to diversity was not supported. The better oxygenated CA seeps at 500 in shared more of the background margin fauna (at 500 m) than did the OR seeps at 800 m (with OMZ fauna at 800 in). Geographical differences in the fluxes of methane-rich fluids and the increased reliance on chemosynthetic food sources with increased depth could explain these results.

2009
Middelburg, JJ, Levin LA.  2009.  Coastal hypoxia and sediment biogeochemistry. Biogeosciences. 6:1273-1293. AbstractWebsite

The intensity, duration and frequency of coastal hypoxia (oxygen concentration < 63 mu M) are increasing due to human alteration of coastal ecosystems and changes in oceanographic conditions due to global warming. Here we provide a concise review of the consequences of coastal hypoxia for sediment biogeochemistry. Changes in bottom-water oxygen levels have consequences for early diagenetic pathways (more anaerobic at expense of aerobic pathways), the efficiency of re-oxidation of reduced metabolites and the nature, direction and magnitude of sediment-water exchange fluxes. Hypoxia may also lead to more organic matter accumulation and burial and the organic matter eventually buried is also of higher quality, i.e. less degraded. Bottom-water oxygen levels also affect the organisms involved in organic matter processing with the contribution of metazoans decreasing as oxygen levels drop. Hypoxia has a significant effect on benthic animals with the consequences that ecosystem functions related to macrofauna such as bio-irrigation and bioturbation are significantly affected by hypoxia as well. Since many microbes and microbial-mediated biogeochemical processes depend on animal-induced transport processes (e.g. re-oxidation of particulate reduced sulphur and denitrification), there are indirect hypoxia effects on biogeochemistry via the benthos. Severe long-lasting hypoxia and anoxia may result in the accumulation of reduced compounds in sediments and elimination of macrobenthic communities with the consequences that biogeochemical properties during trajectories of decreasing and increasing oxygen may be different (hysteresis) with consequences for coastal ecosystem dynamics.

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.

Neira, C, Delgadillo-Hinojosa F, Zirino A, Mendoza G, Levin LA, Porrachia M, Deheyn DD.  2009.  Spatial distribution of copper in relation to recreational boating in a California shallow-water basin. Chemistry and Ecology. 25:417-433.   10.1080/02757540903334197   AbstractWebsite

The overall effect of the number of boats on the copper (Cu) levels in the water column and sediment, along with their spatial variability within Shelter Island Yacht Basin (SIYB), San Diego Bay, California was examined. We identified a horizontal gradient of increasing dissolved Cu and Cu in sediment from outside to the head of SIYB which was coincident with the increasing number of boats. Spatial models of Cu distribution in water and sediment indicated the presence of 'hotspots' of Cu concentration. From outside to the head of SIYB, dissolved Cu ranged from 1.3 gL-1 to 14.6 gL-1 in surface water, and 2.0 gL-1 to 10.2 gL-1 in bottom water. Cu in sediment exceeded the Effect Range Low of 34mgkg-1 (i.e. where adverse effects to fauna may occur), with a peak concentration of 442mgkg-1 at the head of the basin. Free Cu++ in surface water was several orders of magnitude higher than in sediment porewater. High-resolution data of Cu species together with probability maps presented in this paper will allow managers to easily visualise and localise areas of impaired quality and to prioritise which areas should be targeted to improve Cu-related conditions.

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.

2004
Helly, JJ, Levin LA.  2004.  Global distribution of naturally occurring marine hypoxia on continental margins. Deep-Sea Research Part I-Oceanographic Research Papers. 51:1159-1168.   10.1016/j.dsr.2004.03.009   AbstractWebsite

Hypoxia in the ocean influences biogeochemical cycling of elements, the distribution of marine species and the economic well being of many coastal countries. Previous delineations of hypoxic environments focus on those in enclosed seas where hypoxia may be exacerbated by anthropogenically induced eutrophication. Permanently hypoxic water masses in the open ocean, referred to as oxygen minimum zones, impinge on a much larger seafloor surface area along continental margins of the eastern Pacific, Indian and western Atlantic Oceans. We provide the first global quantification of naturally hypoxic continental margin floor by determining upper and lower oxygen minimum zone depth boundaries from hydrographic data and computing the area between the isobaths using seafloor topography. This approach reveals that there are over one million km(2) of permanently hypoxic shelf and bathyal sea floor, where dissolved oxygen is <0.5ml l(-1); over half (59%) occurs in the northern Indian Ocean. We also document strong variation in the intensity, vertical position and thickness of the OMZ as a function of latitude in the eastern Pacific Ocean and as a function of longitude in the northern Indian Ocean. Seafloor OMZs are regions of low biodiversity and are inhospitable to most commercially valuable marine resources, but support a fascinating array of protozoan and metazoan adaptations to hypoxic conditions. (C) 2004 Elsevier Ltd. All rights reserved.

2002
Levin, LA, Michener RH.  2002.  Isotopic evidence for chemosynthesis-based nutrition of macrobenthos: The lightness of being at Pacific methane seeps. Limnology and Oceanography. 47:1336-1345. AbstractWebsite

The importance of chemosynthetic nutritional pathways was examined for macrofaunal invertebrates (>300 mum) from methane seeps in the Gulf of Alaska (4,413-4,443 m), on the Oregon margin (590 m), and on the northern California slope [Eel River margin] (520 m) by use of natural abundance stable isotopic data. Seep macrofauna exhibited lighter delta(13)C and delta(15)N values than those in nonseep sediments, but isotopic signatures varied among seep sites. Macrofaunal isotopic signatures indicated chemosynthetically fixed carbon sources with a significant contribution from methane-derived carbon (MDC) in macrofauna from sediments of pogonophoran fields (average delta(13)C, -46.44parts per thousand, 32%-51% MDC) and Calyptogena phaseoliformis beds (average delta(13)C, -40.89parts per thousand, 12%-40% MDC) in the Gulf of Alaska and in microbial mat sediments on the Oregon margin (average delta(13)C, -43.80parts per thousand, 20%-44% MDC). Lesser influence of MDC was noted in macrofauna from sediments of Calyptogena pacifica beds on the Oregon (average delta(13)C, -33.38parts per thousand, 0%-27% MDC) and California (delta(13)C, -25.10parts per thousand, 0%-22% MDC) margins and from California microbial mat sediments (delta(13)C, -22.23%o, 0%-5% MDC). Although most macrofauna appeared to be heterotrophic, light delta(15)N and delta(13)C values together provided evidence for chemoautotrophic symbioses in selected taxa. Carbon isotopic signatures were consistent with consumption of methane-oxidizing archaea by some dorvilleid polychaetes (delta(13)C, -90.62parts per thousand and -73.80parts per thousand) and with grazing on filamentous sulfur bacteria by gastropods and polychaetes from the Oregon and California seeps. The importance of chemosynthetic trophic pathways varies regionally and among microhabitats, taxonomic groups, and feeding guilds.

Shankle, AM, Goericke R, Franks PJS, Levin LA.  2002.  Chlorin distribution and degradation in sediments within and below the Arabian Sea oxygen minimum zone. Deep-Sea Research Part I-Oceanographic Research Papers. 49:953-969.   10.1016/s0967-0637(01)00077-2   AbstractWebsite

The concentration of chlorophylla degradation products, i.e. chlorins, preserved in deep-sea sediments is a function of the amount of primary production input and the rate at which it is subsequently degraded. Sedimentary chlorins can be used as a proxy for paleoproductivity; however, our understanding of the factors controlling their preservation is limited. To study the effects of changes in export of primary production from the euphotic zone and of differences in depositional conditions on chlorin concentration in marine sediments, chlorins were analyzed by high pressure liquid chromatography from sediments taken within and below the oxygen minimum zone on the Oman margin in the Arabian Sea. Among five stations at water depths between 400 and 1250 m, variation in chlorin concentration in surface sediments (0-0.5 cm) was significantly related to water depth (used here as a proxy for chlorin fluxes to the sediments) and bottom-water oxygen concentration; the more important control on chlorin concentration of surficial sediments measured in this study is the amount of chlorins reaching the sediment. Chlorins decayed exponentially downcore (0-20 cm). The degradation of sedimentary chlorins was better described by a model in which chlorins decayed at different rates within and below the sediment mixed layer. The degradation rates within the mixed layer were 0.0280 +/- 0.0385 yr(-1) (t(1/2) = 73 yr). Below the mixed layer, degradation rates were one to two orders of magnitude less, ranging from 0.0022 +/- 0.0025 yr(-1) (t(1/2) = 680 yr). Many stations had subsurface chlorin concentration peaks between 6 and 10 cm depth. The most likely explanation for these peaks is a period of increased deposition of chlorins in the past. This could result from changes in local depositional environment or a more general increase in surface production resulting in an increased sedimentation of chlorins to the sediments 500-1000 years ago. Chlorins are a useful indicator of the magnitude of chlorin deposition; however their usage as indicators of paleoproductivity is more complicated. (C) 2002 Elsevier Science Ltd. All rights reserved.

2001
Levin, LA, Etter RJ, Rex MA, Gooday AJ, Smith CR, Pineda J, Stuart CT, Hessler RR, Pawson D.  2001.  Environmental influences on regional deep-sea species diversity. Annual Review of Ecology and Systematics. 32:51-93.   10.1146/annurev.ecolsys.32.081501.114002   AbstractWebsite

Most of our knowledge of biodiversity and its causes in the deep-sea benthos derives from regional-scale sampling studies of the macrofauna. Improved sampling methods and the expansion of investigations into a wide variety of habitats have revolutionized our understanding of the deep sea. Local species diversity shows clear geographic variation on spatial scales of 100-1000 km. Recent sampling programs have revealed unexpected complexity in community structure at the landscape level that is associated with large-scale oceanographic processes and their environmental consequences. We review the relationships between variation in local species diversity and the regional-scale phenomena of boundary constraints, gradients of productivity, sediment heterogeneity, oxygen availability, hydrodynamic regimes, and catastrophic physical disturbance. We present a conceptual model of how these interdependent environmental factors shape regional-scale variation in local diversity. Local communities in the deep sea may be composed of species that exist as metapopulations whose regional distribution depends on a balance among global-scale, landscape-scale, and small-scale dynamics. Environmental gradients may form geographic patterns of diversity by influencing local processes such as predation, resource partitioning, competitive exclusion, and facilitation that determine species coexistence. The measurement of deep-sea species diversity remains a vital issue in comparing geographic patterns and evaluating their potential causes. Recent assessments of diversity using species accumulation curves with randomly pooled samples confirm the often-disputed claim that the deep sea supports higher diversity than the continental shelf. However, more intensive quantitative sampling is required to fully characterize the diversity of deep-sea sediments, the most extensive habitat on Earth. Once considered to be constant, spatially uniform, and isolated, deep-sea sediments are now recognized as a dynamic, richly textured environment that is inextricably linked to the global biosphere. Regional studies of the last two decades provide the empirical background necessary to formulate and test specific hypotheses of causality by controlled sampling designs and experimental approaches.

Talley, TS, Levin LA.  2001.  Modification of sediments and macrofauna by an invasive marsh plant. Biological Invasions. 3:51-68.   10.1023/A:1011453003168   AbstractWebsite

Invasive grasses have recently altered salt marsh ecosystems throughout the northern hemisphere. On the eastern seaboard of the USA, Phragmites australis has invaded both brackish and salt marsh habitats. Phragmites australis influence on sediments and fauna was investigated along a salinity and invasion-age gradient in marshes of the lower Connecticut River estuary. Typical salinities were about 19-4 ppt in Site I, 9-10 ppt in Site II and 5-7 ppt in Site III. Strongest effects were evident in the least saline settings (II and III) where Phragmites has been present the longest and exists in monoculture. Limited influence was evident in the most saline region (I) where Phragmites and native salt marsh plants co-occur. The vegetation within Phragmites stands in tidal regions of the Connecticut River generally exhibits taller, but less dense shoots, higher above-ground biomass, and lower below-ground biomass than does the un-invaded marsh flora. There were lower sediment organic content, greater litter accumulation and higher sediment chlorophyll a concentrations in Phragmites-invaded than un-invaded marsh habitat. Epifaunal gastropods (Succinea wilsoni and Stagnicola catascopium) were less abundant in habitats where Phragmites had invaded than in un-invaded marsh habitat. Macro-infaunal densities were lower in the Phragmites-invaded than un-invaded habitats at the two least saline sites (II and III). Phragmites stands supported more podurid insects, sabellid polychaetes, and peracarid crustaceans, fewer arachnids, midges, tubificid and enchytraeid oligochaetes, and greater habitat-wide taxon richness as measured by rarefaction, than did the uninvaded stands. The magnitude and significance of the compositional differences varied with season and with site; differences were generally greatest at the oldest, least saline sites (II and III) and during May, when faunal densities were higher than in September. However, experimental design and the 1-year study period precluded clear separation of salinity, age, and seasonal effects. Although structural effects of Phragmites on salt marsh faunas are evident, further investigation is required to determine the consequences of these effects for ecosystem function.

1994
Levin, LA, Leithold EL, Gross TF, Huggett CL, Dibacco C.  1994.  Contrasting effects of substrate mobility on infaunal assemblages inhabiting two high-energy settings on Fieberling Guyot. Journal of Marine Research. 52:489-522.   10.1357/0022240943077028   AbstractWebsite

The influence of seamount-intensified flows on the structure of infaunal assemblages was examined at two sand-covered sites located 2.3 km apart atop the summit plain of Fieberling Guyot (32-degrees 27.6'N 127-degrees 48.0'W). Both sites experience strong, tidal bottom currents with flows exceeding 20 cm/ s on a daily basis (4 mab). Estimates of shear velocity (u*) did not differ significantly between the two sites. However, differences in sediment composition and density produced different sediment transport regimes at the two sites. At Sea Pen Rim (SPR), located on the NW perimeter (635 m), sedimentary particles were composed primarily of basaltic sands that experienced negligible transport during the study period. At White Sand Swale (WSS, 580 m), a narrow valley enclosed on three sides by basalt outcrops, sediments were composed almost entirely of foraminiferal sands that moved daily. Sediment organic content and microbial abundances were similar at the two sites. Infauna (> 300 mum) had higher densities at WSS (1870/m2) than SPR (1489/m2), but lower expected species richness. Although the 2 sites shared nearly 50% of identified species, peracarid crustaceans, echinoderms, sponges, and bryozoans were proportionally more important in the stable substrates of SPR, while turbellarians, bivalves, and aplacophorans were better represented in the shifting sands of WSS. The infauna of WSS lived deeper in the sediment column (> 50% below 2 cm) than that of SPR (> 50% in the upper 1 cm), at least partly because the majority (83%) at WSS were subsurface burrowers with motile lifestyles. Tube-building and epifaunal lifestyles were more common at SPR than WSS, as were surface-deposit and filter-feeding modes. Fences and weirs were deployed at the study sites for 6.5-wk and 6-mo periods to manipulate bottom stress. Changes in faunal patterns within weirs at WSS reinforced our conjecture that contrasting sediment transport regimes explain between-site differences in community structure. Fence effects varied with deployment period and site. Topographic features on Fieberling Guyot produce heterogeneous sedimentary settings characterized by different transport regimes. Our results suggest that substrate mobility exerts primary control over infaunal community structure at the two high-energy sites.

1993
Levin, LA, Huggett D, Myers P, Bridges T, Weaver J.  1993.  Rare earth tagging methods for the study oflarval dispersal by marine invertebrates. Limnology and Oceanography. 38:346-360. AbstractWebsite

Rare-earth elements (REEs) are proposed as a label for invertebrate larvae (clams, barnacles, and polychaetes) and passive larval mimics (polystyrene spheres) to be used in field studies of larval dispersal. Different elements, their uptake levels, retention times, and effects on larval survivorship were examined to assess suitability of REEs as larval tags. Lu, Sm, and Eu were selected for labeling work based on extremely low levels in natural plankton and seawater and due to the high sensitivity of neutron activation analysis to these elements. Adequate label retention occurs on time scales of hours to days in both larvae and microspheres. Combined use of Eu-labeled microspheres and Lu- and Sm-labeled larvae in release-recapture mode offers a highly sensitive method for studying trajectories and behavior of larvae in natural settings. However, dilution factors may limit experimentation to small and intermediate spatial scales (meters to kilometers) in the field.