Publications

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2015
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.

2013
Neira, C, King I, Mendoza G, Sellanes J, De Ley P, Levin LA.  2013.  Nematode community structure along a central Chile margin transect influenced by the oxygen minimum zone. Deep Sea Research Part I: Oceanographic Research Papers. 78:1-15.   http://dx.doi.org/10.1016/j.dsr.2013.04.002   AbstractWebsite

Nematodes are among the metazoans most tolerant of low-oxygen conditions and play major roles in seafloor ecosystem processes. Nematode communities were studied in sediments off Concepción, Central Chile, spanning the outer shelf within the OMZ (122 m) to the mid-lower continental slope (972 m) beneath the OMZ. The total density and biomass of nematodes (core depth 0–10 cm) ranged from 677 to 2006 ind. 10 cm−2, and 168.4 to 506.5 µg DW 10 cm−2, respectively. Among metazoan meiofaunal taxa, nematodes predominated at all sites both in terms of relative abundance (83.7–99.4%) and biomass (53.8–88.1%), followed by copepods, nauplii and polychaetes. Nematodes were represented by 33 genera distributed among 17 families, with densities greatest at low oxygen sites (122–364 m; ~2000 ind. 10 cm−2). Nematode generic and trophic diversity, and individual biomass were lowest, and Rank 1 dominance was highest, at the most oxygen-depleted site (122 m), despite the fact that the organic carbon content of the sediment was maximal at this depth. At the most oxygenated slope sites (827 and 972 m), all of Wieser's nematode feeding groups were represented. In contrast, at the lowest-oxygen site, only selective deposit (bacterial) feeders (1A) were present, indicating a reduction in trophic complexity. A large percentage of nematodes inhabited subsurface sediment layers (>1 cm). At deeper, more oxygenated sites (827 and 972 m), nematode individual biomass increased downcore, while within the OMZ, nematode biomass was low and remained relatively uniform through the sediment column. The concentration of nematodes in deeper sediment layers, the vertical distribution of the feeding groups, as well as the high nutritional quality of the deeper layers, suggest a differential resource partitioning of the food available, which may reduce interspecific competition.

2010
Gooday, AJ, Bett BJ, Escobar E, Ingole B, Levin LA, Neira C, Raman AV, Sellanes J.  2010.  Habitat heterogeneity and its influence on benthic biodiversity in oxygen minimum zones. Marine Ecology-an Evolutionary Perspective. 31:125-147.   10.1111/j.1439-0485.2009.00348.x   AbstractWebsite

Oxygen minimum zones (OMZs; midwater regions with O(2) concentrations <0.5 ml l(-1)) are mid-water features that intercept continental margins at bathyal depths (100-1000 m). They are particularly well developed in the Eastern Pacific Ocean, the Arabian Sea and the Bay of Bengal. Based on analyses of data from these regions, we consider (i) how benthic habitat heterogeneity is manifested within OMZs, (ii) which aspects of this heterogeneity exert the greatest influence on alpha and beta diversity within particular OMZs and (iii) how heterogeneity associated with OMZs influences regional (gamma) diversity on continental margins. Sources of sea-floor habitat heterogeneity within OMZs include bottom-water oxygen and sulphide gradients, substratum characteristics, bacterial mats, and variations in the organic matter content of the sediment and pH. On some margins, hard grounds, formed of phosphorites, carbonates or biotic substrata, represent distinct subhabitats colonized by encrusting faunas. Most of the heterogeneity associated with OMZs, however, is created by strong sea-floor oxygen gradients, reinforced by changes in sediment characteristics and organic matter content. For the Pakistan margin, combining these parameters revealed clear environmental and faunal differences between the OMZ core and the upper and lower boundary regions. In all Pacific and Arabian Sea OMZs examined, oxygen appears to be the master driver of alpha and beta diversity in all benthic faunal groups for which data exist, as well as macrofaunal assemblage composition, particularly in the OMZ core. However, other factors, notably organic matter quantity and quality and sediment characteristics, come into play as oxygen concentrations begin to rise. The influence of OMZs on meiofaunal, macrofaunal and megafaunal regional (gamma) diversity is difficult to assess. Hypoxia is associated with a reduction in species richness in all benthic faunal groups, but there is also evidence for endemism in OMZ settings. We conclude that, on balance, OMZs probably enhance regional diversity, particularly in taxa such as Foraminifera, which are more tolerant of hypoxia than others. Over evolutionary timescales, they may promote speciation by creating strong gradients in selective pressures and barriers to gene flow.

2009
Hughes, DJ, Lamont PA, Levin LA, Packer M, Feeley K, Gage JD.  2009.  Macrofaunal communities and sediment structure across the Pakistan margin Oxygen Minimum Zone, North-East Arabian Sea. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 56:434-448.   10.1016/j.dsr2.2008.05.030   AbstractWebsite

Benthic macrofauna and sediment column features were sampled at five stations along a bathymetric transect (depths 140, 300, 940, 1200, 1850 m) through the Pakistan margin Oxygen Minimum Zone (OMZ) during the 2003 intermonsoon (March-May) and late-post-monsoon (August-October) periods. Objectives were to compare patterns with those described from other OMZs, particularly the Oman margin of the Arabian Sea, in order to assess the relative influence of bottom-water oxygenation and sediment organic content on macrofaunal standing stock and community structure. Macrofaunal density was highest at the 140-m station subject to monsoon-driven shoaling of the OMZ, but there was no elevation of density at the lower OMZ boundary (1200 m). Numbers was extremely low in the OMZ core (300 m) and were not readily explicable from the environmental data. There was no consistent depth-related trend in macrofaunal biomass. Macrofaunal densities were consistently lower than found off Oman but there was less contrast in biomass. A significant post-monsoon decline in macrofaunal density at 140 m was driven by selective loss of polychaete taxa. Polychaeta was the most abundant major taxon at all stations but did not dominate the macrofaunal community to the extent reported from Oman. Cirratulidae and Spionidae were major components of the polychaete fauna at most stations but Acrocirridae, Ampharetidae, Amphinomidae and Cossuridae were more important at 940 m. Polychaete assemblages at each station were almost completely distinct at the species level. Polychaete species richness was positively correlated with bottom-water dissolved oxygen and negatively correlated with sediment TOC, C:N ratio and total phytopigments. Community dominance showed the opposite pattern. The strongly inverse correlation between oxygen and measures of sediment organic content made it difficult to distinguish their relative effects. The strongly laminated sediments in the OMZ core contrasted with the homogeneous, heavily bioturbated sediments above and below this zone but were associated with minimal macrofaunal biomass rather than distinctive functional group composition. In general, data from the Oman margin were weak predictors of patterns seen off Pakistan, and results suggest the importance of local factors superimposed on the broader trends of macrofaunal community composition in OMZs. (C) 2008 Elsevier Ltd. All rights reserved.