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Sellanes, J, Neira C, Quiroga E, Teixido N.  2010.  Diversity patterns along and across the Chilean margin: a continental slope encompassing oxygen gradients and methane seep benthic habitats. Marine Ecology-an Evolutionary Perspective. 31:111-124.   10.1111/j.1439-0485.2009.00332.x   AbstractWebsite

In the present study we review datasets available for the Chilean margin to assess the relationship between environmental (or habitat) heterogeneity and benthic diversity. Several factors, such as the presence of different water masses, including the oxygen-deficient Equatorial Sub-surface Waters (ESSW) at the continental shelf and upper slope, and the Antarctic Intermediate Waters (AIW) at mid slope depths appear to control the bathymetric distribution of benthic communities. The presence of methane seeps and an extended oxygen minimum zone (OMZ) add complexity to the benthic distribution patterns observed. All these factors generate environmental heterogeneity, which is predicted to affect the diversity patterns both along and across the Chilean continental margin. The response to these factors differs among different faunal size groups: meio-, macro-, and megafauna. Physiological adaptations to oxygen deficiency and constraints related to body size of each group seem to explain the larger-scale patterns observed, while sediment/habitat heterogeneity (e.g. at water mass boundaries, hardgrounds, biogeochemical patchiness, sediment organic content, grain size) may influence the local fauna diversity patterns.

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.

Thurber, AR, Kroger K, Neira C, Wiklund H, Levin LA.  2010.  Stable isotope signatures and methane use by New Zealand cold seep benthos. Marine Geology. 272:260-269.   10.1016/j.margeo.2009.06.001   AbstractWebsite

The carbon isotopic composition of seep faunal tissue represents a time-integrated view of the interaction between biology and the biogeochemical gradients within the environment. Here we provide an initial description of carbon and nitrogen stable isotope signatures of dominant symbiont-bearing megafauna and heterotrophic mega- and macrofauna from 10 methane-seep sites on the continental margin of the North Island of New Zealand (662-1201 m water depth). Isotopic signatures suggest that sulfide oxidation supports symbiont-bearing taxa including solemyid and vesicomyid bivalves, and methanotrophic symbionts are present in the seep mussel Bathymodiolus sp Multiple species of Frenulata (Siboglinidae) are present and have a range of isotopic values that are indicative of both thiotroph- and methanotroph-based nutrition. Isotopic composition of the tubeworm Lamellibrachia sp. varied by 23 3 parts per thousand among individuals although there was no consistent difference among sites Variation in methane use by heterotrophic fauna appears to reflect the availability of hard vs. soft substrate, macrofauna on hard substrates had high delta(13)C signatures, reflecting consumption of photosynthetic-derived organic matter Two unique, biotic assemblages were discovered to be fueled largely by methane: a hard-substrate, multi-phyla sponge-associated community (inhabiting the sponge Pseudosuberites sp) and a soft-sediment assemblage dominated by ampharetid polychaetes Isotope signatures yield estimates of 38-100% and 6-100% methane-derived carbon in sponge associates and ampharetid-bed macrofauna. respectively. These estimates are comparable to those made for deeper methane seeps at the Florida Escarpment (3290 m) and Kodiak. Alaska seeps (4445 m) The overall high use of methane as a carbon source by both symbiont-bearing and heterotrophic fauna suggests that New Zealand methane seeps are an ideal model system to study the interaction among metazoans, bacteria, archaea, and their resulting effect on methane cycles. (C) 2009 Elsevier B V All rights reserved

Rathburn, AE, Levin LA, Tryon M, Gieskes JM, Martin JM, Perez ME, Fodrie FJ, Neira C, Fryer GJ, Mendoza G, McMillan PA, Kluesner J, Adamic J, Ziebis W.  2009.  Geological and biological heterogeneity of the Aleutian margin (1965-4822 m). Progress in Oceanography. 80:22-50.   10.1016/j.pocean.2008.12.002   AbstractWebsite

Geological, biological and biogeochemical characterization of the previously unexplored margin off Unimak Island, Alaska between 1965 and 4822 m water depth was conducted to examine: (1) the geological processes that shaped the margin, (2) the linkages between depth, geomorphology and environmental disturbance in structuring benthic communities of varying size classes and (3) the existence, composition and nutritional sources of methane seep biota on this margin. The study area was mapped and sampled using multibeam sonar, a remotely operated vehicle (ROV) and a towed camera system. Our results provide the first characterization of the Aleutian margin mid and lower slope benthic communities (micro-biota, foraminifera, macrofauna and megafauna), recognizing diverse habitats in a variety of settings. Our investigations also revealed that the geologic feature known as the "Ugamak Slide" is not a slide at all, and could not have resulted from a large 1946 earthquake. However, sediment disturbance appears to be a pervasive feature of this margin. We speculate that the deep-sea occurrence of high densities of Elphidium, typically a shallow-water foraminiferan, results from the influence of sediment redeposition from shallower habitats. Strong representation of cumacean, amphipod and tanaid crustaceans among the Unimak macrofauna may also reflect sediment instability. Although some faunal abundances decline with depth, habitat heterogeneity and disturbance generated by canyons and methane seepage appear to influence abundances of biota in ways that supercede any clear depth gradient in organic matter input. Measures of sediment organic matter and pigment content as well as C and N isotopic signatures were highly heterogeneous, although the availability of organic matter and the abundance of microorganisms in the upper sediment (1-5 cm) were positively correlated. We report the first methane seep on the Aleutian slope in the Unimak region (3263-3285 m), comprised of clam bed, pogonophoran field and carbonate habitats. Seep foraminiferal assemblages were dominated by agglutinated taxa, except for habitats above the seafloor on pogonophoran tubes. Numerous infaunal taxa in clam bed and pogonophoran field sediments and deep-sea "reef' cnidarians (e.g., corals and hydroids) residing on rocks near seepage sites exhibited light organic delta(13)C signatures indicative of chemosynthetic nutritional sources. The extensive geological, biogeochemical and biological heterogeneity as well as disturbance features observed on the Aleutian slope provide an attractive explanation for the exceptionally high biodiversity characteristic of the world's continental margins. (C) 2008 Elsevier Ltd. All rights reserved.

Levin, L, Gutierrez D, Rathburn A, Neira C, Sellanes J, Munoz P, Gallardo V, Salamanca M.  2002.  Benthic processes on the Peru margin: a transect across the oxygen minimum zone during the 1997-98 El Nino. Progress in Oceanography. 53:1-27.   10.1016/s0079-6611(02)00022-8   AbstractWebsite

Oxygen minimum zones (OMZs) are widespread features in the most productive regions of the world ocean. A holistic view of benthic responses to OMZ conditions will improve our ability to predict ecosystem-level consequences of climatic trends that influence oxygen availability, such as global warming or ENSO-related events. Four stations off Callao, Peru (-12'S, Station A, 305 m; Station B, 562 m; Station C, 830 nu and Station D, 1210 m) were sampled to examine the influence of the low bottom-water oxygen concentration and high organic-matter availability within the OMZ (O(2) < 0.5 ml L(-1)) on sediments, benthic communities, and bioturbation. Sampling took place during early January 1998, an intense El Ni (n) over tildeo period associated with higher-than-normal levels of O(2) on the shelf and upper slope. Peru slope sediments were highly heterogeneous. Sediment total organic carbon content exceeded 16%, lamination was present below 6 cm depth, and filamentous sulfur bacteria (Thioploca spp.) were present at Station A, (305 m, 0, < 0.02 ml L(-1)). Deeper sites contained phosphorite crusts or pellets and exhibited greater bottom-water oxygenation and lower content and quality of organic matter. X-radiographs and (210)Pb and (234)Th profiles suggested the dominance of lateral transport and bioturbation over pelagic sedimentation at the mid- and lower slope sites. Macrofauna, metazoan meiofauna and foraminifera exhibited coherence of density patterns across stations, with maximal densities (and for macrofauna, reduced diversity) at Station A, where bottom-water oxygen concentration was lowest and sediment labile organic matter content (LOC: sum of protein, carbohydrate and lipid carbon) was greatest. Metazoan and protozoan meiofaunal densities were positively correlated with sediment LOC. The taxa most tolerant of nearly anoxic, organic-rich conditions within the Peru OMZ were calcareous foraminifera, nematodes and gutless phallodrilinid (symbiont-bearing) oligochaetes. Agglutinated foraminifera, harpacticoid copepods, polychaetes and many other macrofaunal taxa increased in relative abundance below the OMZ. During the study (midpoint of the 1997-98 El Ni (n) over tildeo), the upper OMZ boundary exhibited a significant deepening (to 190 m) relative to 'normal', non-El Ni (n) over tildeo conditions (< 100 m), possibly causing a mild, transient oxygenation over the upper slope (200-300 m) and reduction of the organic particle flux to the seabed. Future sampling may determine whether the Peru margin system exhibits dynamic responses to changing ENSO-related conditions. (C) 2002 Elsevier Science Ltd. All rights reserved.

Neira, C, Sellanes J, Levin LA, Arntz WE.  2001.  Meiofaunal distributions on the Peru margin: relationship to oxygen and organic matter availability. Deep-Sea Research Part I-Oceanographic Research Papers. 48:2453-2472.   10.1016/s0967-0637(01)00018-8   AbstractWebsite

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12 degreesS). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440-548 ind. 10cm(-2) (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2-5 cm. interval. At the deeper sites, about 50% were restricted to the top I cm. The importance of copepods and nauplii increased consistently with depth, reaching similar to 12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02mll(-1) supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r = 0.99, p <0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic. heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites. (C) 2001 Elsevier Science Ltd. All rights reserved.

Neira, C, Rackemann M.  1996.  Black spots produced by buried macroalgae in intertidal sandy sediments of the Wadden Sea: Effects on the meiobenthos. Journal of Sea Research. 36:153-170.   10.1016/s1385-1101(96)90786-8   AbstractWebsite

The effects of buried decaying macroalgae on meiobenthos were examined in intertidal sandy sediments of the Wadden Sea of Lower Saxony. In situ experiments confirmed that one of the principal causes of the formation of reduced surface sediments or 'black spots' on the tidal flats is the increasing occurrence and subsequent decomposition of filamentous green algae (Enteromorpha spp.) buried in the sediment. Five to fifteen days after algal material had been buried, the sediment surface turned black. The impact of these black spots on meiobenthos was dramatic: the changed chemical conditions in the sediment resulted in long and drastic reductions in meiofaunal abundance and number of taxa. A multi-dimensional scaling (MDS) analysis of data on meiobenthic abundances revealed that samples from black-spot areas were clearly separated from those of control and reference areas. Re-oxidized black spots showed recolonization by meiofaunal animals, with numbers of individuals and taxa similar to those of oxidized surface sediments. The use of abundances of members of higher meiobenthic taxa to monitor changes in the sediment's chemistry, especially those caused by biomass overload, is discussed.