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Neira, C, Ingels J, Mendoza G, Hernandez-Lopez E, Levin LA.  2018.  Distribution of meiofauna in bathyal sediments influenced by the oxygen minimum zone off Costa Rica. Frontiers in Marine Science. 5   10.3389/fmars.2018.00448   AbstractWebsite

Ocean deoxygenation has become a topic of increasing concern because of its potential impacts on marine ecosystems, including oxygen minimum zone (OMZ) expansion and subsequent benthic effects. We investigated the influence of oxygen concentration and organic matter (OM) availability on metazoan meiofauna within and below an OMZ in bathyal sediments off Costa Rica, testing the hypothesis that oxygen and OM levels are reflected in meiofaunal community structures and distribution. Mean total densities in our sampling cores (400-1800 m water depth) were highest with 3688 ind. 10 cm(-2) at the OMZ core at 400 m water depth, decreasing rapidly downslope. Nematodes were overall dominant, with a maximum of 99.9% in the OMZ core, followed by copepods (13%), nauplii (4.8%), and polychaetes (3%). Relative copepod and nauplii abundance increased consistently with depth and increasing bottom-water O-2. Meiofaunal composition was significantly different among sites, with lower taxonomic diversity at OMZ sites relative to deeper, oxygenated sites. Vertical distribution patterns within sediments showed that in strongly oxygen-depleted sites less meiofauna was concentrated in the surface sediment than at deeper slope sites. Highest meiofaunal abundance and lowest diversity occurred under lowest oxygen and highest pigment levels, whereas highest diversity occurred under highest oxygen-concentrations and low pigments, as well as high quality of sedimentary pigment (chl a/phaeo) and organic carbon (C/N). The lower meiofaunal diversity, and lower structural and trophic complexity, at oxygen-depleted sites raises concerns about changes in the structure and function of benthic marine ecosystems in the face of OMZ expansions.

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.

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.

Sellanes, J, Quiroga E, Neira C, Gutierrez D.  2007.  Changes of macrobenthos composition under different ENSO cycle conditions on the continental shelf off central Chile. Continental Shelf Research. 27:1002-1016.   10.1016/j.csr.2007.01.001   AbstractWebsite

The course of environmental conditions and shelf macrobenthic communities off Central Chile (similar to 36 degrees S) during the strong 1997-98 El Nino (EN) event is compared with a subsequent and basically "normal" period (2002-2003). Changes in macrofaunal community, feeding mode structure, and biomass size spectra are contrasted over time with changes in oceanographic and sediment settings, in order to assess intra- and inter-annual changes in faunal composition during both ENSO periods. During EN, there was a decrease in biomass and abundance of species known to be well adapted to organic-rich, oxygen-deficient environments, such as the interface-feeding polychacte Paraprionospio pinnata. On the other hand the abundance of highly mobile, burrowing polychaetes remained unaffected, or even increased in biomass. The decline of P. pinnata lasted several years after the demise of warm conditions, possibly due to negative interactions with those more mobile burrowing polychaetes. The percent contribution of subsurface-deposit feeders to total biomass increased during EN (49.3 +/- 12.4% during summer) and declined only in the summer-fall period of 2002-03 (11.1 +/- 4.1%). An opposite trend was observed for interface and surface-deposit feeders. From EN to summer-fall 2002-03 (i.e., normal to low oxygen conditions) the size-structure of the macrobenthos switched from a larger to a smaller-sized assemblage. However, biomass was maintained due to successful recruitment and high abundance of both P. pinnata and the squat lobster, Pleuroncodes monodon. Our results suggest that the shelf macrofaunal community structure exhibit fluctuations at various time scales, and that these changes are more pronounced during and after a strong EN event. In such cases, the effects of such an event may be recorded at latitudes as far south as 36 degrees S, with consequences in the biota lasting for many years after the demise of warm conditions. (c) 2007 Elsevier Ltd. All rights reserved.

Contreras, S, Pantoja S, Neira C, Lange CB.  2007.  Biogeochemistry of surface sediments off Concepcion (~36°S), Chile: El Nino vs. non-El Nino conditions. Progress in Oceanography. 75:576-585.   10.1016/j.pocean.2007.08.030   AbstractWebsite

We compared the signals of several water column properties (upwelling intensity, sea level anomaly, temperature, nutrients, dissolved oxygen, chlorophyll-a, and surface sediments) of the continental shelf off Concepcion (36 degrees S) during the 1997-1998 El Nino with those of a normal year (2002-2003). We found that the primary hydrographic effect of El Nino 1997-1998 was a reduction in the input of nutrient-rich, oxygen-poor Equatorial Subsurface Water over the shelf. This affected the biology of the water column, as evidenced by the reduced phytoplankton biomass. Surface sediment properties (biogenic opal, organic carbon, bulk delta N-15) observed during El Nino 1997-1998 reflected a reduced export production and the sediments failed to show the water column seasonality that occurs under normal conditions. In addition, weakened denitrification and/or upper water column fertilization could be inferred from the sedimentary delta N-15. Although diminished, export production was preserved in the surface sediments, revealing less degraded organic matter in the upwelling period of the El Nino year than in the normal year. We suggest that the fresher organic material on the seafloor was probably associated with a severe reduction in the polychaete Parapronospio pinnata, which is considered to be the most important metazoan remineralizer of organic carbon at the sediment-water interface in the study area. (c) 2007 Elsevier Ltd. All rights reserved.

Sellanes, J, Neira C.  2006.  ENSO as a natural experiment to understand environmental control of meiofaunal community structure. Marine Ecology-an Evolutionary Perspective. 27:31-43.   10.1111/j.1439-0485.2005.00069.x   AbstractWebsite

The sediments of the Bay of Concepcion and the adjacent shelf underlie one of the most productive upwelling areas in the SE Pacific margin. Reports on factors controlling meiofaunal community structure in these kinds of organic-rich and oxygen-deficient habitats are scarce in the literature. In this study, five sites along a transect from the mid-Bay of Concepcion (27 m) to the outer shelf (120 m) were studied on fives dates (May, August, November 1997, and March and May 1998) in order to assess the dynamic relationships between sedimentary organic matter and metazoan meiofauna. The sampling period coincided with the 1997-1998 El Nino event. Sediment parameters investigated were the redox potential discontinuity depth, photosynthetic pigment concentrations (chlorophyll a and phaeopigments), organic carbon, nitrogen, total lipids, carbohydrates, and proteins. In general, lowest values of meiofauna abundance and biomass were found within the naturally eutrophic Bay of Concepcion and towards the shelf break, while maximum values occurred at intermediate depths. During the whole period, the meiofaunal abundance was negatively correlated with the concentration of most of the biochemical components of organic matter, as well as with the sediment phaeopigment content. However, positive correlations were found with chlorophyll a derived indices and with bottom-water oxygen content. Most of the sediment parameters displayed a seasonal cycle, but towards the beginning of 1998, an effect of the 1997-1998 El Nino was evident. Typical austral -summer (i.e. oxygen-deficient) conditions did not develop, and sedimentary parameters reflected a decreased input of phytodetritus. Along the transect, the magnitude of this effect on meiofauna varied among sites. An overall positive response, in terms of meiofaunal abundance was observed, probably due to the amelioration of low oxygen conditions in the sediment.

Levin, LA, Neira C, Grosholz ED.  2006.  Invasive cordgrass modifies wetland trophic function. Ecology. 87:419-432.   10.1890/04-1752   AbstractWebsite

Vascular plants strongly control belowground environments in most ecosystems. Invasion by vascular plants in coastal wetlands, and by cordgrasses (Spartina spp.) in particular, are increasing in incidence globally, with dramatic ecosystem-level consequences. We examined the trophic consequences of' invasion by a Spartina hybrid (S. alterniflora X S. foliosa) in San Francisco Bay (USA) by documenting differences in biomass and trophic structure of benthic communities between sediments invaded by Spartina and uninvaded sediments. We found the invaded system shifted from all algae-bascd to a detritus-based food web. We then tested for a relationship between diet and tolerance to invasion, hypothesizing that species that consume Spartina detritus are more likely to inhabit invaded sediments than those that consume surface algae. Infaunal diets were initially examined with natural abundance stable isotope analyses and application of mixing models, but these yielded an ambiguous picture of food sources. Therefore, we conducted isotopic enrichment experiments by providing N-15-labeled Spartina detritus both on and below the sediment surface in areas that either contained Spartina or were unvegetated. Capitellid and nereid polychaetes, and oligochaetes, groups shown to persist following Spartina invasion of San Francisco Bay tidal flats, took up N-15 from labeled native and invasive Spartina detritus. In contrast, We found that amphipods, bivalves, and other taxa less tolerant to invasion consumed primarily surficial algae, based oil C-13 enrichment experiments. Habitat (Spartina vs. unvegetated patches) and location of' detritus (on or within sediments) did not affect N-15 uptake from cletritus. Our investigations support a "trophic shift" model for ecosystem response to wetland plant invasion and preview loss of key trophic support for fishes and migratory birds by shifting dominance to species not widely consumed by species at higher trophic levels.

Neira, C, Decraemer W, Backeljau T.  2005.  A new species of Glochinema (Epsilonematidae : Nematoda) from the oxygen minimum zone off Baja California, NE Pacific and phylogenetic relationships at species level within the family. Cahiers De Biologie Marine. 46:105-126. AbstractWebsite

Glochinema spinithorni sp. nov. is described from muddy, bathyal sediments of the oxygen minimum zone (OMZ) of the north eastern Pacific Ocean off Baja California. It is characterized by a rather large body length (over I mm), a large number of body rings (242-282), sexual dimorphism in the number and position of pharyngeal thorns near the head region, cuticular ornamentation with numerous hairy spines and two sets of stronger spines ventrally in the anterior body half, mid-sized spiral amphids, and by the number and arrangement of ambulatory setae and long spicules (over 100 mu m). The finding of a second OMZ Glochinema species with similar morphological characteristics supports the presumption that OMZs function as isolated habitats promoting endemism at low taxonomic levels. Results of phylogenetic analyses at the species level restricted to the Glochinematinae and Keratonematinae suggest that both OMZ species form a monophyletic group. An attempt was made to recognize phylogenetic relationships at species level within the family. The phylogenetic analyses were based upon a data matrix of 96 ingroup taxa and 25 characters. The results support the monophyly of the genera Leptepsilonema, Metepsilonema, Perepsilonema and Polkepsilonema (with or without Pternepsilonema).

Munoz, P, Salamanca MA, Neira C, Sellanes J.  2004.  Nitrogen sediment fluxes in an upwelling system off central Chile (Concepcion Bay and adjacent shelf) during the 1997-1998 El Nino. Revista Chilena De Historia Natural. 77:305-318.   10.4067S0716-078X2004000200009   AbstractWebsite

Two stations representing coastal and shelf environments (bay of Concepcion and adjacent shelf, respectively) were sampled in order to assess benthic nitrogen fluxes. During the study period anomalously high oxygen bottom waters conditions were observed, coinciding with the 1997-1998 El Nino event. From both sites, intact sediment cores were collected for incubation experiments to directly estimate total ammonium fluxes. Pore-water chemistry was also assessed to estimate diffusive fluxes of nitrogen compounds (NH(4)(+), NO(3)(-) and NO(2)(-)), The NH(4)(+) flux from incubation experiments ranged from 0.55 to 2.18 mmol m(-2) day(-1) in the inner bay and from 1.84 to 2.14 mmol m(-2) day(-1) on the shelf. NO(3)(-) diffusive fluxes to the sediments were observed in the bay in March (-17 x 10(-2) mmol m(-2) day(-1)) and on the shelf in November (-2.7 x 10(-2) mmol m(-2) day(-1)), associated with the lowest NH(4)(+) diffusive fluxes and coincident with a stronger gradient of Eh in the first centimeters of the sediments. Besides, total NH(4)(+) fluxes lower than in normal periods were observed, suggesting the influence of less reducing conditions, consequence in turn of the lower fluxes of particulated organic material and higher dissolved oxygen in the bottom waters. These conditions were probably imposed by the El Nino event. On the other hand, and relative to previous reports, an increase of irrigation coefficients (DI) was estimated. Thus, the flux extension of other nitrogen compounds could have acquired relevance in relation to NH(4)(+) fluxes during the study period.

Sellanes, J, Neira C, Quiroga E.  2003.  Composition, structure and energy flux of the meiobenthos off central Chile. Revista Chilena De Historia Natural. 76:401-415.   10.4067S0716-078X2003000300006.   AbstractWebsite

The general objective of this study was to determine the structure of metazoan meiofauna (at a high taxonomic level) and to estimate its role in the energy flux of the benthic sub-system off Concepcion, Chile (similar to36degrees30' S). Samples were collected in May and November 1997 and May 1998 at five sites located at the: inner Bay of Concepcion (28 m), bay-mouth (35 m), inner-shelf (64 m), mid-shelf (88 in) and outer-shelf (120 m). The study period coincided with the El Nino 1997-1998 (EN) event. The diversity of the meiobenthos was low, but density and biomass were moderate to high compared with average values reported for muddy sub-littoral environments elsewhere. Nematodes were the dominant group, with over 95 % (10(3)-10(4) ind 10 cm(-2)) of the total density, followed by copepods and polychaetes. Highest densities and biomasses were always found at the bay-mouth, followed by the inner-bay and the inner-shelf, while the lowest values occurred at the mid- and outer-shelf. Off Concepcion, the meiofauna plays an important role in the energy flux through the benthic sub-system. Indeed, the meiofauna could be converting to biomass and remineralizing in the inner-bay and bay-mouth up to 36 and 45 %, respectively, of the organic carbon reaching the sediments from the water column.

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.