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2017
Hansman, RL, Thurber AR, Levin LA, Aluwihare LI.  2017.  Methane fates in the benthos and water column at cold seep sites along the continental margin of Central and North America. Deep-Sea Research Part I-Oceanographic Research Papers. 120:122-131.   10.1016/j.dsr.2016.12.016   AbstractWebsite

The potential influence of methane seeps on carbon cycling is a key question for global assessments, but the study of carbon cycling in surface sediments and the water column of cold seep environments is complicated by the high temporal and spatial variability of fluid and gas fluxes at these sites. In this study we directly examined carbon sources supporting benthic and planktonic food webs at venting methane seeps using isotopic and molecular approaches that integrate this variability. At four seep environments located along North and Central America, microorganisms from two size fractions were collected over several days from 2800 to 90501 of seawater to provide a time-integrated measure of key microbial groups and the carbon sources supporting the overall planktonic microbial community. In addition to water column measurements, the extent of seafloor methane release was estimated at two of the sites by examining the stable carbon isotopic signature (delta C-13) of benthic metazoan infauna. This signature reveals carbon sources fueling the base of the food chain and thus provides a metric that represents a time-integrated view of the dominant microbial processes within the sediment. The stable carbon isotopic composition of microbial DNA (delta C-13-DNA), which had values between -17.0 and -19.5%(0), indicated that bulk planktonic microbial production was not ultimately linked to methane or other C-13-depleted seep-derived carbon sources. Instead these data support the importance of organic carbon derived from either photo- or chemoautotrophic CO2 fixation to the planktonic food web. Results of qPCR of microbial DNA sequences coding for a subunit of the particulate methane monooxygenase gene (pmoA) showed that only a small percentage of the planktonic microbial community were potential methane oxidizers possessing pmoA (< 5% of 16S rRNA gene copies). There was an overall decrease of C-13-depleted carbon fueling the benthic metazoan community from 3 to 5 cm below the seafloor to the sediment surface, reflecting limited use of isotopically depleted carbon at the sediment surface. Rare methane emission as indicated by limited aerobic methane oxidation acts to corroborate our findings for the planktonic microbial community.

2007
Woulds, C, Cowie GL, Levin LA, Andersson JH, Middelburg JJ, Vandewiele S, Lamont PA, Larkin KE, Gooday AJ, Schumacher S, Whitcraft C, Jeffreys RM, Schwartz M.  2007.  Oxygen as a control on seafloor biological communities and their roles in sedimentary carbon cycling. Limnology and Oceanography. 52:1698-1709.   10.4319/lo.2007.52.4.1698   AbstractWebsite

C-13 tracer experiments were conducted at sites spanning the steep oxygen, organic matter, and biological community gradients across the Arabian Sea oxygen minimum zone, in order to quantify the role that benthic fauna play in the short-term processing of organic matter (OM) and to determine how this varies among different environments. Metazoan macrofauna and macrofauna-sized foraminiferans took up as much as 56 +/- 13 mg of added C m(-2) (685 mg C m(-2) added) over 2-5 d, and at some sites this uptake was similar in magnitude to bacterial uptake and/or total respiration. Bottom-water dissolved oxygen concentrations exerted a strong control over metazoan macrofaunal OM processing. At oxygen concentrations > 7 mu mol L-1 (0.16 ml L-1), metazoan macrofauna were able to take advantage of abundant OM and to dominate OM uptake, while OM processing at O-2 concentrations of 5.0 mu mol L-1 (0.11 ml L-1) was dominated instead by (macrofaunal) foraminiferans. This led us to propose the hypothesis that oxygen controls the relative dominance of metazoan macrofauna and foraminifera in a threshold manner, with the threshold lying between 5 and 7 mu mol L-1 (0.11 to 0.16 ml L-1). Large metazoan macrofaunal biomass and high natural concentrations of OM were also associated with rapid processing of fresh OM by the benthic community. Where they were present, the polychaete Linopherus sp. and the calcareous foraminiferan Uvigerina ex gr. semiornata, dominated the uptake of OM above and below, respectively, the proposed threshold concentrations of bottom-water oxygen.

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.

1991
Levin, LA, Childers SE, Smith CR.  1991.  Epibenthic, agglutinating foraminiferans in the Santa Catalina Basin and their response to disturbance. Deep-Sea Research Part a-Oceanographic Research Papers. 38:465-483.   10.1016/0198-0149(91)90047-j   AbstractWebsite

There are five common species of large (0.5-6 cm long) epibenthic, agglutinating foraminiferans in the Santa Catalina Basin (1200-1350 m). This paper describes their basic ecology and response to mound disturbance. Combined, the five species attain mean densities of 200-300 individuals per m2 and their protoplasm has an average biomass of 199.5 mg m-2. Individual species occur at densities ranging from 7 to 100 m-2, and each species has a different population size structure. Protoplasm comprises < 2% of test volumes. Analysis of excess Th-234 revealed no indication of particle sequestering within tests, and acridine orange direct counts of bacteria provided no evidence of microbial gardening or enhancement associated with tests. Twenty-five per cent of tests examined had metazoan associates; approximately half of these were polychaetes. Experiments were carried out to investigate the response of the epibenthic foraminiferal assemblage to disturbance from large, biogenic mounds, a common feature on the Santa Catalina Basin floor. Three branched forms, Pelosina cf. arborescens, P. cf. cylindrica and a mud-walled astrorhizinid, were most abundant on background sediments, less common on natural mounds and absent from artificially-created mounds exposed for 10.5 months. Two spherical species, Oryctoderma sp. and a different mud-walled astrorhizinid, were present at similar densities on artificial mounds (9.5-10.5 months old), natural mounds and undisturbed sediments, but Oryctoderma sp. attained largest sizes on mounds. These two species appear to be opportunistic taxa that can colonize and grow rapidly on mound sediments. This study suggests that disturbance, in this case that by sediment mound builders, is an important source of spatial heterogeneity in deep-water foraminiferal communities. Where sediment mounds occur, foraminiferal assemblages will experience disequilibrium dynamics.