Publications

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2012
Bernardino, AF, Levin LA, Thurber AR, Smith CR.  2012.  Comparative composition, diversity and trophic ecology of sediment macrofauna at vents, seeps and organic falls. Plos One. 7   10.1371/journal.pone.0033515   AbstractWebsite

Sediments associated with hydrothermal venting, methane seepage and large organic falls such as whale, wood and plant detritus create deep-sea networks of soft-sediment habitats fueled, at least in part, by the oxidation of reduced chemicals. Biological studies at deep-sea vents, seeps and organic falls have looked at macrofaunal taxa, but there has yet to be a systematic comparison of the community-level attributes of sediment macrobenthos in various reducing ecosystems. Here we review key similarities and differences in the sediment-dwelling assemblages of each system with the goals of (1) generating a predictive framework for the exploration and study of newly identified reducing habitats, and (2) identifying taxa and communities that overlap across ecosystems. We show that deep-sea seep, vent and organic-fall sediments are highly heterogeneous. They sustain different geochemical and microbial processes that are reflected in a complex mosaic of habitats inhabited by a mixture of specialist (heterotrophic and symbiont-associated) and background fauna. Community-level comparisons reveal that vent, seep and organic-fall macrofauna are very distinct in terms of composition at the family level, although they share many dominant taxa among these highly sulphidic habitats. Stress gradients are good predictors of macrofaunal diversity at some sites, but habitat heterogeneity and facilitation often modify community structure. The biogeochemical differences across ecosystems and within habitats result in wide differences in organic utilization (i.e., food sources) and in the prevalence of chemosynthesis-derived nutrition. In the Pacific, vents, seeps and organic-falls exhibit distinct macrofaunal assemblages at broad-scales contributing to beta diversity. This has important implications for the conservation of reducing ecosystems, which face growing threats from human activities.

2004
Gallardo, VA, Palma M, Carrasco FD, Gutierrez D, Levin LA, Canete JI.  2004.  Macrobenthic zonation caused by the oxygen minimum zone on the shelf and slope off central Chile. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 51:2475-2490.   10.1016/j.dsr2.2004.07.028   AbstractWebsite

The relationship between macrobenthic (greater than or equal to 300 mum) zonation and the oxygen minimum zone (OMZ: O(2) < 0.5 ml L(-1)) was studied in shelf and slope sediments (122-840 m depth) off Concepcion Bay, central Chile. Four study sites were sampled during March-April 1999 for abiotic factors, macrofaunal density, biomass, mean individual size, and diversity. Within the OMZ (122-206 m), the macrofaunal density was high (16,478-21,381 individuals m(-2)) and 69-89% of the organisms were soft-bodied. Density was highest (21,381 individuals m(-2)), biomass lowest (16.95 g wet weight m-2), and individual size smallest (0.07 mg C individuals) at the shelf break site (206 m). Polychaete worms made up 71% of the total abundance, crustaceans 16%, and mollusks only 2%. Total abundance beneath the OMZ (mid-slope site, similar to840 m) was 49% crustaceans and 43% polychaetes. Although existing literature originally led to the hypothesis that both diversity and biomass within the OMZ would be lower than beneath the OMZ, in the present study this was only true for diversity. Biomass distribution, on the other hand, was concave along the depth gradient; the highest values were near the upper edge of (122 m) and beneath (840 m) the OMZ. Indices of the macrofaunal community structure varied in relation to bottom-water oxygen concentration, chlorophyll-alpha, phaeopigments, and sulfide concentration, but not in relation to grain size, C, N, mud, porosity, redox potential, a bottom-water temperature. (C) 2004 Published by Elsevier Ltd.

2000
Gooday, AJ, Bernhard JM, Levin LA, Suhr SB.  2000.  Foraminifera in the Arabian Sea oxygen minimum zone and other oxygen-deficient settings: taxonomic composition, diversity, and relation to metazoan faunas. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 47:25-54.   10.1016/s0967-0645(99)00099-5   AbstractWebsite

Previous work has shown that some foraminiferal species thrive in organically enriched, oxygen-depleted environments. Here, we compare 'live' (stained) faunas in multicorer samples (0-1 cm layer) obtained at two sites on the Oman margin, one located at 412m within the oxygen minimum zone (OMZ) (O(2) = 0.13 ml l(-1)), the other located at 3350 m, well below the main OMZ (O(2) similar to 3.00 ml l(-1)). While earlier studies have focused on the hard-shelled (predominantly calcareous) foraminifera, we consider complete stained assemblages, including poorly known, soft-shelled, monothalamous forms. Densities at the 412-m site were much higher (16,107 individuals.10 cm(-2) in the > 63-mu m fraction) than at the 3350-m site (625 indiv.10 cm(-2)). Species richness (E(S(100))), diversity (H', Fishers Alpha index) and evenness (J') were much lower, and dominance (R1D) was higher, at 412 m compared with 3350 m. At 412 m, small calcareous foraminifera predominated and soft-shelled allogromiids and sacamminids were a minor faunal element. At 3350 m, calcareous individuals were much less common and allogromiids and saccamminids formed a substantial component of the fauna. There were also strong contrasts between the foraminiferal macrofauna( > 300-mu m fraction) at these two sites; relatively small species of Bathysiphon, Globobulimina and Lagenammina dominated at 412 m, very large, tubular, agglutinated species of Bathysiphon, Hyperammina, Rhabdammina and Saccorhiza were important at 3350 m. Our observations suggest that, because they contain fewer soft-shelled and agglutinated foraminifera, a smaller proportion of bathyal, low-oxygen faunas is lost during fossilization compared to faunas from well-oxygenated environments. Trends among foraminifera (> 63 mu m fraction) in the Santa Barbara Basin (590 and 610m depth; O(2) = 0.05 and 0.15 ml(-1) respectively), and macrofaunal foraminifera(> 300 mu m) on the Peru margin (300-1250 m depth: O(2) = 0.02-1.60 mi l(-1)), matched those observed on the Oman margin. Tn particular, soft-shelled monothalamous taxa were rare and large agglutinated taxa were absent in the most oxygen-depleted ( < 0.20 mi l(-1)) stations. Foraminifera often outnumber metazoans (both meiofaunal and macrofaunal) in bathyal oxygen-depleted settings. However, although phylogenetically distant, foraminifera and metazoans exhibit similar population responses to oxygen depletion; species diversity decreases, dominance increases, and the relative abundance of the major taxa changes. The foraminiferal macrofauna ( > 300 mu m) were 5 times more abundant than the metazoan macrofauna at 412 m on the Oman margin but 16 times more abundant at the 3350 m site. Among the meiofauna (63-300 mu m), the trend was reversed, foraminifera were 17 times more abundant than metazoan taxa at 412 m but only 1.4 times more abundant at 3350 In. An abundance of food combined with oxygen levels which are not depressed sufficiently to eliminate the more tolerant taxa, probably explains why foraminifera and macrofaunal metazoans flourished at the 412-m site, perhaps to the detriment of the metazoan meiofauna. (C) 1999 Elsevier Science Ltd. All rights reserved.

Levin, LA, Gage JD, Martin C, Lamont PA.  2000.  Macrobenthic community structure within and beneath the oxygen minimum zone, NW Arabian Sea. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 47:189-226.   10.1016/s0967-0645(99)00103-4   AbstractWebsite

Investigations of macrobenthos were carried out within and beneath the oxygen minimum zone (OMZ, < 0.5 ml l(-1)) during Fall 1994 on the Oman margin, NW Arabian Sea. Six stations (400, 700, 850, 1000, 1250 and 3400m) were characterized with respect to macrofaunal abundance, biomass, body size, taxonomic composition, diversity and lifestyles, and the relation of these parameters to environmental conditions. The OMZ (400-1000 m) was dominated by a dense (5818-19,183 ind m(-2)), soft-bodied assemblage consisting largely (86-99%) of surface-feeding polychaetes, Spionids and cirratulids dominated at the 400- and 700-m stations, paraonids and ampharetids at the 850- and 1000-m stations. Molluscs and most crustaceans were common only below the OMZ ( greater than or equal to 1250 m); a species of the amphipod Ampelisca was abundant within the OMZ, however. Both density and biomass were elevated within the OMZ relative to stations below but body size did not differ significantly among stations. The lower OMZ boundary (0.5 ml l(-1)) was not a zone of enhanced macrofaunal standing stock, as originally hypothesized. However, abundance maxima at 700-850m may reflect an oxygen threshold (0.15-0.20 ml l(-1)) above which macrofauna take advantage of organically enriched sediments. Incidence of burrowing and subsurface-deposit feeding increased below the OMZ, Species richness (E[S(100)]), diversity (H') and evenness (J') were lower and dominance (R1D) was higher within than beneath the OMZ. Within-station (between-boxcore) faunal heterogeneity increased markedly below the OMZ. Surface sediment pigment concentrations and oxygen together explained 96-99% of the variance in measures of E[S(100)], H' and J' across the transect; grain size and % TOC did not yield significant regressions. Pigments, assumed to reflect food availability and possibly oxygen effects on organic matter preservation, were negatively correlated with species richness and evenness, and positively correlated with dominance. The reverse was true for water depth. Macrobenthic patterns of calcification and lifestyle within the Oman margin OMZ (0.13-0.3 mi l(-1)) match the dysaerobic biofacies of paleo-environmental reconstruction models. (C) 1999 Elsevier Science Ltd. All rights reserved.

1994
Levin, LA.  1994.  Paleoecology and ecology of xenophyophores. Palaios. 9:32-41.   10.2307/3515076   AbstractWebsite

Xenophyophores are large (several mm to 25 cm diameter), agglutinating protozoans, found primarily in the deep sea. Tests range from simple fans, discs or mudballs, to elaborately folded or reticulated forms, and often contain specific particle types or sizes. Xenophyophore densities are highest on sloped sediments associated with seamounts, continental slopes, canyons and trenches, and beneath productive surface waters. Most forms live as epibenthos on hard or soft substrates, feeding on surface deposits and suspended particles, but one endobenthic genus has been described. Modern xenophyophores enhance particle flux to the seabed, creating local regions of intense radiotracer and metazoan activity. Features key to xenophyophore identification in the fossil record include distinct test morphologies that sometimes involve incorporation of globigerinacean tests, sandwich-like structure in cross section, concentrations of barite (found within the protoplasm), and the presence of fecal strands containing many 10 to 20 mm fecal pellets having enhanced Pb contents. While xenophyophores have no confirmed fossil record, modern structures resembling the ichnogenus Paleodictyon are made by endobenthic xenophyophores in the genus Occultammina. Parallel distributions, morphologies and behaviors have been proposed for some graphoglyptid trace fossils, and for some xenophyophores in the families Syringamminidae and Reticulamminidae. Both occur in the deep sea, have regular or irregular network morphologies, and have been proposed to garden bacteria, trap meiofauna, or suspension feed. Fossils previously regarded as phylloid and fucoid algae, and several species of Aschemonella, previously regarded as foraminiferids, have lifestyles and morphologies consistent with those of modern epibenthic xenophyophores. Confirmation of xenophyophore presence in stratigraphic sequences could provide paleohabitat information and help elucidate the origins of this protozoan group.