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Neira, C, Levin LA, Mendoza G, Zirino A.  2014.  Alteration of benthic communities associated with copper contamination linked to boat moorings. Marine Ecology-an Evolutionary Perspective. 35:46-66.   10.1111/maec.12054   AbstractWebsite

Although copper (Cu) is an essential element for life, leaching from boat paint can cause excess environmental loading in enclosed marinas. The effects of copper contamination on benthic macrofaunal communities were examined in three San Diego Bay marinas (America's Cup, Harbor Island West and East) in Southern California, USA. The distribution of Cu concentration in sediments exhibited a clear spatial gradient, with hotspots created by the presence of boats, which in two marinas exceeded the effect range medium (ERM). Elevated sediment Cu was associated with differences in benthic assemblages, reduced species richness and enhanced dominance in America's Cup and Harbor Island West, whereas Harbor Island East did not appear to be affected. At sites without boats there were greater abundances of some amphipods such as the species Desdimelita sp., Harpinia sp., Aoroides sp., Corophium sp., Podocerus sp., bivalves such as Lyonsia californica, Musculista senhousia, Macoma sp., and polychaetes such as Diplocirrus sp. In contrast, at sites with boats, densities of Pseudopolydora paucibranchiata, Polydora nuchalis, Euchone limnicola, Exogone lourei, Tubificoides spp. were enhanced. The limited impact on Harbor Island East suggests not only lower Cu input rates and increased water flushing and mixing, but also the presence of adequate defense mechanisms that regulate availability and mitigate toxic impacts. At all three marinas, Cu in tissues of several macrobenthic species exhibited Cu bioaccumulation above levels found in the surrounding environment. The annelids Lumbrineris sp. and Tubificoides spp., and the amphipod Desdimelita sp. contained high levels of Cu, suggesting they function as Cu bioaccumulators. The spionid polychaetes Polydora nuchalis and Pseudopolydora paucibranchiata had much lower Cu concentrations than surrounding sediments, suggesting they function as Cu bioregulators. The macrobenthic invertebrates in San Diego Bay marinas that tolerate Cu pollution (e.g. P.nuchalis, P.paucibranchiata, Euchone limnicola, Typosyllis sp., Tubificoides sp.) may function as indicators of high-Cu conditions, whereas the presence of Cu-sensitive species (e.g. Podocerus sp., Aoroides sp., Harpinia sp., Macoma sp., Lyonsia californica) may indicate healthier conditions (less Cu-stressed). Parallel responses by faunas of Shelter Island Yacht Basin, also in San Diego Bay, suggest potential for development of regional Cu contamination assessment criteria, and call for functional comparisons with other marinas and coastal water bodies.

Marlow, JJ, Steele JA, Ziebis W, Thurber AR, Levin LA, Orphan VJ.  2014.  Carbonate-hosted methanotrophy represents an unrecognized methane sink in the deep sea. Nature Communications. 5   10.1038/ncomms6094   AbstractWebsite

The atmospheric flux of methane from the oceans is largely mitigated through microbially mediated sulphate-coupled methane oxidation, resulting in the precipitation of authigenic carbonates. Deep-sea carbonates are common around active and palaeo-methane seepage, and have primarily been viewed as passive recorders of methane oxidation; their role as active and unique microbial habitats capable of continued methane consumption has not been examined. Here we show that seep-associated carbonates harbour active microbial communities, serving as dynamic methane sinks. Microbial aggregate abundance within the carbonate interior exceeds that of seep sediments, and molecular diversity surveys reveal methanotrophic communities within protolithic nodules and well-lithified carbonate pavements. Aggregations of microbial cells within the carbonate matrix actively oxidize methane as indicated by stable isotope FISH-nanoSIMS experiments and (CH4)-C-14 radiotracer rate measurements. Carbonate-hosted methanotrophy extends the known ecological niche of these important methane consumers and represents a previously unrecognized methane sink that warrants consideration in global methane budgets.

Zirino, A, Neira C, Maicu F, Levin LA.  2013.  Comments on and implications of a steady-state in coastal marine ecosystems. Chemistry & Ecology. 29:86-99.   10.1080/02757540.2012.696613   AbstractWebsite

Coastal ecosystems can be thought of as being established by a number of physico-geochemical drivers, e.g. geochemistry and bathymetry of the basins, climate, tidal and freshwater flows, natural and anthropogenic inputs of nutrients and toxins, all of which exert an influence on the resulting communities of organisms. Depending on the interactions among the major drivers, ecosystems may occur on both large and small scales and be basin-wide or within basins. For individual and separate ecosystems to exist with some permanence in time, e.g. reach a steady-state, they also have to be ‘defended’. Defences are mechanisms that counter changes to maintain the status quo. We argue, and present evidence to support the notion, that the defence mechanisms are inextricably tied to primary production and the biogeochemical cycling of organic matter and provide buffers that mitigate potentially adverse impacts by trace toxins. Colloid pumping, production of complexing ligands and sulfide formation are some of the mechanisms that control trace substances. Current methods for assessing ecosystems do not address the issue of steady-state, nor do they take account of defence activities, e.g. buffering. Therefore, they cannot assess the ‘robustness’ of ecosystems or their ability to resist change, for good or bad. Also, defence mechanisms may, for a time, mask future potentially serious impacts, suggesting that monitoring efforts with limited budgets should consider the measurement of the inputs into ecosystems as well as the immediate or short-term result of the inputs. [ABSTRACT FROM PUBLISHER]Copyright of Chemistry & Ecology is the property of Taylor & Francis Ltd and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Breitburg, D, Levin LA, Oschlies A, Grégoire M, Chavez FP, Conley DJ, Garçon V, Gilbert D, Gutiérrez D, Isensee K, Jacinto GS, Limburg KE, Montes I, Naqvi SWA, Pitcher GC, Rabalais NN, Roman MR, Rose KA, Seibel BA, Telszewski M, Yasuhara M, Zhang J.  2018.  Declining oxygen in the global ocean and coastal waters. Science. 359   10.1126/science.aam7240   Abstract

As plastic waste pollutes the oceans and fish stocks decline, unseen below the surface another problem grows: deoxygenation. Breitburg et al. review the evidence for the downward trajectory of oxygen levels in increasing areas of the open ocean and coastal waters. Rising nutrient loads coupled with climate change—each resulting from human activities—are changing ocean biogeochemistry and increasing oxygen consumption. This results in destabilization of sediments and fundamental shifts in the availability of key nutrients. In the short term, some compensatory effects may result in improvements in local fisheries, such as in cases where stocks are squeezed between the surface and elevated oxygen minimum zones. In the longer term, these conditions are unsustainable and may result in ecosystem collapses, which ultimately will cause societal and economic harm.

Moseman, SM, Zhang R, Qian PY, Levin LA.  2009.  Diversity and functional responses of nitrogen-fixing microbes to three wetland invasions. Biological Invasions. 11:225-239.   10.1007/s10530-008-9227-0   AbstractWebsite

Impacts of invasive species on microbial components of wetland ecosystems can reveal insights regarding functional consequences of biological invasions. Nitrogen fixation (acetylene reduction) rates and diversity of nitrogen fixers, determined by genetic fingerprinting (T-RFLP) of the nifH gene, were compared between native and invaded sediments in three systems. Variable responses of nitrogen fixing microbes to invasion by a non-native mussel, Musculista senhousia, and mangrove, Avicennia marina, in Kendall Frost-Northern Wildlife Preserve (Mission Bay) and salt cedar, Tamarisk (Tamarix spp.) in Tijuana Estuary suggest microbes respond to both species- and site-specific influences. Structurally similar invaders (the mangrove and salt cedar) produced different effects on activity and diversity of nitrogen fixers, reflecting distinct environmental contexts. Despite relative robustness of microbial community composition, subtle differences in total diversity or activity of nitrogen fixers reveal that microbes are not immune to impacts of biological invasions, and that functional redundancy of microbial diversity is limited, with significant consequences for functional dynamics of wetlands.

Rabalais, NN, Diaz RJ, Levin LA, Turner RE, Gilbert D, Zhang J.  2010.  Dynamics and distribution of natural and human-caused hypoxia. Biogeosciences. 7:585-619. AbstractWebsite

Water masses can become undersaturated with oxygen when natural processes alone or in combination with anthropogenic processes produce enough organic carbon that is aerobically decomposed faster than the rate of oxygen re-aeration. The dominant natural processes usually involved are photosynthetic carbon production and microbial respiration. The re-supply rate is indirectly related to its isolation from the surface layer. Hypoxic water masses (< 2 mg L(-1), or approximately 30% saturation) can form, therefore, under 'natural' conditions, and are more likely to occur in marine systems when the water residence time is extended, water exchange and ventilation are minimal, stratification occurs, and where carbon production and export to the bottom layer are relatively high. Hypoxia has occurred through geological time and naturally occurs in oxygen minimum zones, deep basins, eastern boundary upwelling systems, and fjords. Hypoxia development and continuation in many areas of the world's coastal ocean is accelerated by human activities, especially where nutrient loading increased in the Anthropocene. This higher loading set in motion a cascading set of events related to eutrophication. The formation of hypoxic areas has been exacerbated by any combination of interactions that increase primary production and accumulation of organic carbon leading to increased respiratory demand for oxygen below a seasonal or permanent pycnocline. Nutrient loading is likely to increase further as population growth and resource intensification rises, especially with increased dependency on crops using fertilizers, burning of fossil fuels, urbanization, and waste water generation. It is likely that the occurrence and persistence of hypoxia will be even more widespread and have more impacts than presently observed. Global climate change will further complicate the causative factors in both natural and human-caused hypoxia. The likelihood of strengthened stratification alone, from increased surface water temperature as the global climate warms, is sufficient to worsen hypoxia where it currently exists and facilitate its formation in additional waters. Increased precipitation that increases freshwater discharge and flux of nutrients will result in increased primary production in the receiving waters up to a point. The interplay of increased nutrients and stratification where they occur will aggravate and accelerate hypoxia. Changes in wind fields may expand oxygen minimum zones onto more continental shelf areas. On the other hand, not all regions will experience increased precipitation, some oceanic water temperatures may decrease as currents shift, and frequency and severity of tropical storms may increase and temporarily disrupt hypoxia more often. The consequences of global warming and climate change are effectively uncontrollable at least in the near term. On the other hand, the consequences of eutrophication-induced hypoxia can be reversed if long-term, broad-scale, and persistent efforts to reduce substantial nutrient loads are developed and implemented. In the face of globally expanding hypoxia, there is a need for water and resource managers to act now to reduce nutrient loads to maintain, at least, the current status.

Levin, LA, Ziebis W, Mendoza GF, Bertics VJ, Washington T, Gonzalez J, Thurber AR, Ebbed B, Lee RW.  2013.  Ecological release and niche partitioning under stress: Lessons from dorvilleid polychaetes in sulfidic sediments at methane seeps. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 92:214-233.   10.1016/j.dsr2.2013.02.006   AbstractWebsite

Organisms inhabiting methane seep sediments are exposed to stress in the form of high levels of hydrogen sulfide, which result mainly from sulfate reduction coupled to anaerobic methane oxidation. Dorvilleidae (Polychaeta) have successfully invaded this ecosystem, and multiple species in divergent genetic clades co-occur at high densities. At methane seeps in the NE Pacific off California and Oregon, the genera Ophryotrocha, Parougia and Exallopus are especially well represented. To test the hypothesis that dorvilleid coexistence is facilitated by niche partitioning through sulfide tolerance and trophic patterns, we examined dorvilleid species-specific patterns of occurrence and nutrition at methane seeps off Eel R. [ER] on the Californian continental slope and at Hydrate Ridge [HR] on the Oregon continental slope, and in two habitats (clam bed and microbial mat) characterized by lower and higher hydrogen sulfide levels, respectively. Microelectrode measurements of hydrogen sulfide enabled characterization of environmental sulfide levels for species sampled in background sediment cores and in colonization trays. Dorvilleids tolerated H2S levels from 10 mu M to over 2.6 mM, with the majority of species inhabiting sediments with similar environmental H2S concentrations (median 85-100 mu M). Dorvilleid species richness was greater at HR than ER, but did not differ between clam bed and microbial mat habitats. Species distribution patterns reflected preferences for ER clam bed (lower sulfide levels), ER mat and HR clam bed (moderate sulfide levels), or HR mat (very high sulfide levels). Nutritional patterns, including trophic diversity and functional similarity, were examined using community stable isotope metrics based on delta N-15 and delta C-13. Within each region, dorvilleid species exhibited multiple trophic strategies. Co-existing congeners typically exhibited distinct isotope signatures, suggesting trophic partitioning. Trophic diversity and delta N-15 range for whole assemblages (measured by Total Hull Area and Standard Elliptical Area using species averages) and functional redundancy or species packing (measured as distance to nearest neighbor) among species and individuals were generally higher at ER, where sulfide levels were lower than at HR. In contrast, average trophic diversity among individuals within a species was greater at HR than ER. In colonization experiments involving agar-based manipulations of sulfide in tray sediments that mimicked clam bed and mat conditions, dorvilleids comprised 68% and 48% of colonists at ER and HR, respectively. Dorvilleid species richness was higher in trays that were initially more sulfidic. However, habitat exerted stronger influence on the composition of colonizing dorvilleids than did sulfide additions. In the NE Pacific, regional, habitat and vertical (down-core) variation in hydrogen sulfide creates complex environmental heterogeneity at methane seeps, promoting high diversity of stress-tolerant taxa such as dorvilleid polychaetes. (C) 2013 Elsevier Ltd. All rights reserved.

Levin, LA, Ekau W, Gooday AJ, Jorissen F, Middelburg JJ, Naqvi SWA, Neira C, Rabalais NN, Zhang J.  2009.  Effects of natural and human-induced hypoxia on coastal benthos. Biogeosciences. 6:2063-2098.   10.5194/bg-6-2063-2009   AbstractWebsite

Coastal hypoxia (defined here as < 1.42 ml L(-1); 62.5 mu M; 2 mg L(-1), approx. 30% oxygen saturation) develops seasonally in many estuaries, fjords, and along open coasts as a result of natural upwelling or from anthropogenic eutrophication induced by riverine nutrient inputs. Permanent hypoxia occurs naturally in some isolated seas and marine basins as well as in open slope oxygen minimum zones. Responses of benthos to hypoxia depend on the duration, predictability, and intensity of oxygen depletion and on whether H(2)S is formed. Under suboxic conditions, large mats of filamentous sulfide oxidizing bacteria cover the seabed and consume sulfide. They are hypothesized to provide a detoxified microhabitat for eukaryotic benthic communities. Calcareous foraminiferans and nematodes are particularly tolerant of low oxygen concentrations and may attain high densities and dominance, often in association with microbial mats. When oxygen is sufficient to support metazoans, small, soft-bodied invertebrates (typically annelids), often with short generation times and elaborate branchial structures, predominate. Large taxa are more sensitive than small taxa to hypoxia. Crustaceans and echinoderms are typically more sensitive to hypoxia, with lower oxygen thresholds, than annelids, sipunculans, molluscs and cnidarians. Mobile fish and shellfish will migrate away from low-oxygen areas. Within a species, early life stages may be more subject to oxygen stress than older life stages. Hypoxia alters both the structure and function of benthic communities, but effects may differ with regional hypoxia history. Human-caused hypoxia is generally linked to eutrophication, and occurs adjacent to watersheds with large populations or agricultural activities. Many occurrences are seasonal, within estuaries, fjords or enclosed seas of the North Atlantic and the NW Pacific Oceans. Benthic faunal responses, elicited at oxygen levels below 2 ml L(-1), typically involve avoidance or mortality of large species and elevated abundances of enrichment opportunists, sometimes prior to population crashes. Areas of low oxygen persist seasonally or continuously beneath upwelling regions, associated with the upper parts of oxygen minimum zones (SE Pacific, W Africa, N Indian Ocean). These have a distribution largely distinct from eutrophic areas and support a resident fauna that is adapted to survive and reproduce at oxygen concentrations < 0.5 ml L(-1). Under both natural and eutrophication-caused hypoxia there is loss of diversity, through attrition of intolerant species and elevated dominance, as well as reductions in body size. These shifts in species composition and diversity yield altered trophic structure, energy flow pathways, and corresponding ecosystem services such as production, organic matter cycling and organic C burial. Increasingly the influences of nature and humans interact to generate or exacerbate hypoxia. A warmer ocean is more stratified, holds less oxygen, and may experience greater advection of oxygen-poor source waters, making new regions subject to hypoxia. Future understanding of benthic responses to hypoxia must be established in the context of global climate change and other human influences such as overfishing, pollution, disease, habitat loss, and species invasions.

Levin, LA, Zhu J, Creed E.  1991.  The genetic basis of life history characters in a polychaete exhibiting planktotrophy and lecithotrophy. Evolution. 45:380-397.   10.2307/2409672   AbstractWebsite

The polychaete Streblospio benedicti is unusual in that several field populations consist of individuals that exhibit either planktotrophic or lecithotrophic larval development. Planktotrophy in this species involves production of many small ova that develop into feeding larvae with a two- to three-week planktonic period. Lecithotrophy involves production of fewer, larger ova that develop into nonfeeding larvae that are brooded longer and have a brief planktonic stage. Reciprocal matings were performed to investigate genetic variance components and the correlation structure of life-history traits associated with planktotrophy and lecithotrophy. Our objective was to better understand persistence of this developmental dichotomy in Streblospio benedicti, and among marine invertebrates in general. Substantial additive genetic variation (75-98% of total) was detected for the following characters at first reproduction: female length; position of the first gametogenic setiger and first brood pouch; ovum diameter; three traits related to fecundity (numbers of ova per ovary, larvae per brood pouch, and larvae per brood); median larval planktonic period and the presence of larval swimming setae; but not for total number of brood pouches; larval length; larval feeding; and larval survivorship. Based on the unusual geographic distribution of development modes in this species, we hypothesize that the developmental traits have evolved in allopatry and have only recently come into contact in North Carolina. The high additive contribution to variance observed for many traits may be inflated due to (a) nonrandom breeding in nature, and (b) examination of only one component of an age-structured population at one time. Nuclear interaction variance and maternal variance accounted for 84% of the total variation in larval survivorship. This observation supports other empirical studies and theoretical predictions that nonadditive components of variance will increase in importance in individual traits that are most closely tied to fitness. A network of life-history trait correlations was observed that defines distinct planktotrophic and lecithotrophic trait complexes. Negative genetic correlations were present between fecundity and egg size, between fecundity and position of the first gametes, and between larval survivorship and median planktonic period. Positive genetic correlations were detected between fecundity and female size at first reproduction and between planktonic period and the presence of swimming setae. Intergenerational product-moment correlations were negative for larval length and fecundity, planktonic period and egg size, female size and larval survivorship, and fecundity and larval survivorship. If the genetic correlation structure observed in the laboratory persists in the field, it may constrain responses of individual characters to directional selection, and indirectly perpetuate the dichotomies associated with planktotrophy and lecithotrophy.

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.

Gooday, AJ, Jorissen F, Levin LA, Middelburg JJ, Naqvi SWA, Rabalais NN, Scranton M, Zhang J.  2009.  Historical records of coastal eutrophication-induced hypoxia. Biogeosciences. 6:1707-1745.   10.5194/bg-6-1707-2009   AbstractWebsite

Under certain conditions, sediment cores from coastal settings subject to hypoxia can yield records of environmental changes over time scales ranging from decades to millennia, sometimes with a resolution of as little as a few years. A variety of biological and geochemical indicators (proxies) derived from such cores have been used to reconstruct the development of eutrophication and hypoxic conditions over time. Those based on (1) the preserved remains of benthic organisms (mainly foraminiferans and ostracods), (2) sedimentary features (e.g. laminations) and (3) sediment chemistry and mineralogy (e.g. presence of sulphides and redox-sensitive trace elements) reflect conditions at or close to the seafloor. Those based on (4) the preserved remains of planktonic organisms (mainly diatoms and dinoflagellates), (5) pigments and lipid biomarkers derived from prokaryotes and eukaryotes and (6) organic C, N and their stable isotope ratios reflect conditions in the water column. However, the interpretation of these indicators is not straightforward. A central difficulty concerns the fact that hypoxia is strongly correlated with, and often induced by, organic enrichment caused by eutrophication, making it difficult to separate the effects of these phenomena in sediment records. The problem is compounded by the enhanced preservation in anoxic and hypoxic sediments of organic microfossils and biomarkers indicating eutrophication. The use of hypoxia-specific proxies, such as the trace metals molybdenum and rhenium and the bacterial biomarker isorenieratene, together with multi-proxy approaches, may provide a way forward. All proxies of bottom-water hypoxia are basically qualitative; their quantification presents a major challenge to which there is currently no satisfactory solution. Finally, it is important to separate the effects of natural ecosystem variability from anthropogenic effects. Despite these problems, in the absence of historical data for dissolved oxygen concentrations, the analysis of sediment cores can provide plausible reconstructions of the temporal development of human-induced hypoxia, and associated eutrophication, in vulnerable coastal environments.

Neira, C, Mendoza G, Levin LA, Zirino A, Delgadillo-Hinojosa F, Porrachia M, Deheyn DD.  2011.  Macrobenthic community response to copper in Shelter Island Yacht Basin, San Diego Bay, California. Marine Pollution Bulletin. 62:701-717.   10.1016/j.marpolbul.2011.01.027   AbstractWebsite

We examined Cu contamination effects on macrobenthic communities and Cu concentration in invertebrates within Shelter Island Yacht Basin, San Diego Bay, California. Results indicate that at some sites, Cu in sediment has exceeded a threshold for "self defense" mechanisms and highlight the potential negative impacts on benthic faunal communities where Cu accumulates and persists in sediments. At sites with elevated Cu levels in sediment, macrobenthic communities were not only less diverse but also their total biomass and body size (individual biomass) were reduced compared to sites with lower Cu. Cu concentration in tissue varied between species and within the same species, reflecting differing abilities to "regulate" their body load. The spatial complexity of Cu effects in a small marina such as SIYB emphasizes that sediment-quality criteria based solely on laboratory experiments should be used with caution, as they do not necessarily reflect the condition at the community and ecosystem levels. (C) 2011 Elsevier Ltd. All rights reserved.

Zhang, J, Gilbert D, Gooday AJ, Levin L, Naqvi SWA, Middelburg JJ, Scranton M, Ekau W, Pena A, Dewitte B.  2010.  Natural and human-induced hypoxia and consequences for coastal areas: synthesis and future development. Biogeosciences. 7:1443-1467., France: European Geosciences Union AbstractWebsite

Hypoxia has become a world-wide phenomenon in the global coastal ocean and causes a deterioration of the structure and function of ecosystems. Based on the collective contributions of members of SCOR Working Group #128, the present study provides an overview of the major aspects of coastal hypoxia in different biogeochemical provinces, including estuaries, coastal waters, upwelling areas, fjords and semi-enclosed basins, with various external forcings, ecosystem responses, feedbacks and potential impact on the sustainability of the fishery and economics. The obvious external forcings include freshwater runoff and other factors contributing to stratification, organic matter and nutrient loadings, as well as exchange between coastal and open ocean water masses. Their different interactions set up mechanisms that drive the system towards hypoxia. Coastal systems also vary in their relative susceptibility to hypoxia depending on their physical and geographic settings. It is understood that coastal hypoxia has a profound impact on the sustainability of ecosystems, which can be seen, for example, by the change in the food-web structure and system function; other influences include compression and loss of habitat, as well as changes in organism life cycles and reproduction. In most cases, the ecosystem responds to the low dissolved oxygen in non-linear ways with pronounced feedbacks to other compartments of the Earth System, including those that affect human society. Our knowledge and previous experiences illustrate that there is a need to develop new observational tools and models to support integrated research of biogeochemical dynamics and ecosystem behavior that will improve confidence in remediation management strategies for coastal hypoxia.

Zapata-Hernandez, G, Sellanes J, Thurber AR, Levin LA, Chazalon F, Linke P.  2014.  New insights on the trophic ecology of bathyal communities from the methane seep area off Concepcion, Chile (similar to 36 degrees S). Marine Ecology-an Evolutionary Perspective. 35:1-21.   10.1111/maec.12051   AbstractWebsite

Studies of the trophic structure in methane-seep habitats provide insight into the ecological function of deep-sea ecosystems. Methane seep biota on the Chilean margin likely represent a novel biogeographic province; however, little is known about the ecology of the seep fauna and particularly their trophic support. The present study, using natural abundance stable isotopes, reveals a complex trophic structure among heterotrophic consumers, with four trophic levels supported by a diversity of food sources at a methane seep area off Concepcion, Chile (similar to 36 degrees S). Although methanotrophy, thiotrophy and phototrophy are all identified as carbon fixation mechanisms fueling the food web within this area, most of the analysed species (87.5%) incorporate carbon derived from photosynthesis and a smaller number (12%) use carbon derived from chemosynthesis. Methane-derived carbon (MDC) incorporation was documented in 22 taxa, including sipunculids, gastropods, polychaetes and echinoderms. In addition, wide trophic niches were detected in suspension-feeding and deposit-feeding taxa, possibly associated with the use of organic matter in different stages of degradation (e.g. from fresh to refractory). Estimates of Bayesian standard ellipses area (SEA(B)) reveal different isotopic niche breadth in the predator fishes, the Patagonian toothfish Dissostichus eleginoides and the combtooth dogfish Centroscyllium nigrum, suggesting generalist versus specialist feeding behaviors, respectively. Top predators in the ecosystem were the Patagonian toothfish D. eleginoides and the dusky cat shark, Bythaelurus canescens. The blue hake Antimora rostrata also provides a trophic link between the benthic and pelagic systems, with a diet based primarily on pelagic-derived carrion. These findings can inform accurate ecosystem models, which are critical for effective management and conservation of methane seep and adjacent deep-sea habitats in the Southeastern Pacific.

Thorrold, SR, Zacherl DC, Levin LA.  2007.  Population connectivity and larval dispersal using geochemical signatures in calcified structures. Oceanography. 20:80-89.   dx.doi.org/10.5670/oceanog.2007.31   AbstractWebsite

The importance of larval dispersal to the population dynamics and biogeography of marine organisms has been recognized for almost a century (Hjort, 1914; Thorson, 1950). More recently, theoretical studies have highlighted the role that connectivity may play in determining the resilience of marine populations (Hastings and Botsford, 2006). Effective spatial management of marine capture fisheries, including the design of marine reserve networks, also requires an understanding of population connectivity (Sale et al., 2005). However, remarkably few empirical estimates of larval dispersal or population connectivity in ocean environments exist.

Levin, LA, Ziebis W, Mendoza GF, Growney-Cannon V, Walther S.  2006.  Recruitment response of methane-seep macrofauna to sulfide-rich sediments: An in situ experiment. Journal of Experimental Marine Biology and Ecology. 330:132-150.   10.1016/j.jembe.2005.12.022   AbstractWebsite

Hydrodynamically unbiased colonization trays were deployed for 6 months (Oct. 2000 to April 2001) on the northern California margin (Eel R. region; 525 m) to examine macrofaunal colonization rates at methane seeps. The influence of sulfide on recruitment and survival was examined by deploying sediments with and without sulfide added; effect of seep proximity was evaluated by placing trays inside and outside seeps. The trays contained a two-layer system mimicking vesicomyid clam bed habitat geochemistry, with 89 9 mM sulfide in a lower agar layer at the start of the experiment. After 6 month on the seabed, the lower agar layer contained 2-4 mM H2S. We observed rapid macrofaunal colonization equivalent to 50% of initial non-seep ambient densities. There was no difference in total colonizer densities, number of species, or rarefaction diversity among 3 treatments: (1) controls (no sulfide added) placed outside seeps, (2) trays with sulfide added placed outside seeps and (3) trays with sulfide added placed inside seep patches. Colonization trays with sulfide placed at seeps had different species composition from trays without sulfide place outside seeps; there were more amphipods (non-ampeliscid) and cumaceans in the seep/sulfide treatment and more nemerteans, Nephtys cornuta and tanaids in the non-seep/no-sulfide treatment. Outside seeps, annelids comprised <15% of tray colonists; within seep patches, annelids comprised 5 of the top 10 dominant colonizing taxa (24% of the total). The polychaetes Mediomastus sp., Aphelochaeta sp., Paraonidae sp., and Nerillidae sp. exhibited significantly higher densities in sulfide additions. Tanaids, echinoderms, and N. cornuta exhibited sulfide avoidance. At least 6 dorvilleid polychaete species colonized the experiments. Of these, 4 species occurred exclusively in trays with sulfide added and 80% of all dorvilleid individuals were found in trays with sulfide placed inside seep sediments. Counts of large sulfur bacterial filaments were positively correlated with maximum sulfide concentration in each tray, and with proximity of sulfide to the sediment surface. However, total macrofaunal densities were not correlated with tray sulfide concentrations. As a group, tray assemblages achieved some but not all characteristics of ambient seep assemblages after 6-month exposure on the sea floor. Distinctive colonization patterns at methane seeps contribute to the dynamic mosaic of habitat patches that characterize the eastern Pacific continental margin. Overall, proximity of seep habitats had at least as great an influence on macrofaunal colonization as tray sulfide concentrations. Taxa characteristic of seep sediments were more likely to settle into trays placed inside rather than outside seep patches. Whether this is due to limited dispersal ability or local geochemical cues remains to be determined. (C) 2005 Elsevier B.V. All rights reserved.

Zirino, A, Elwany H, Neira C, Maicu F, Mendoza G, Levin LA.  2014.  Salinity and its variability in the Lagoon of Venice, 2000–2009. Advances in Oceanography and Limnology. 5:41-59.: Taylor & Francis   10.1080/19475721.2014.900113   Abstract

Yearly averages computed from monthly and bimonthly salinity data collected between 2000 and 2009 from 13 broadly spaced stations in the Venice Lagoon were analysed in view of 30 min data collected semi-continuously during 2009 at nine similarly located stations. Data from all stations and all years indicate that, based on yearly averages, the lagoon may be divided along its major (long) axis into three areas: 1) a northern, freshwater impacted area (S = <28 PSU) of high, tidally-caused, variability, 2) a southern, marine, zone of S >32 PSU of low, tidally-caused, variability, and 3) an intermediate zone. Salinity changes are closely associated with rainfall events, and the incoming freshwater is consistently distributed throughout the lagoon by tidal action. Much variability is simply a result of the forward and backward motion of the tides and is not caused by a salinity change in the water itself. The consistency of the 2000?2009 data and the historical (to 1961) watershed record support the hypothesis that the Venice Lagoon has been and is currently at steady-state with respect to its salinity distribution. As such, it is conducive to the development of (at least) three separate ecosystems.

Neira, C, Delgadillo-Hinojosa F, Zirino A, Mendoza G, Levin LA, Porrachia M, Deheyn DD.  2009.  Spatial distribution of copper in relation to recreational boating in a California shallow-water basin. Chemistry and Ecology. 25:417-433.   10.1080/02757540903334197   AbstractWebsite

The overall effect of the number of boats on the copper (Cu) levels in the water column and sediment, along with their spatial variability within Shelter Island Yacht Basin (SIYB), San Diego Bay, California was examined. We identified a horizontal gradient of increasing dissolved Cu and Cu in sediment from outside to the head of SIYB which was coincident with the increasing number of boats. Spatial models of Cu distribution in water and sediment indicated the presence of 'hotspots' of Cu concentration. From outside to the head of SIYB, dissolved Cu ranged from 1.3 gL-1 to 14.6 gL-1 in surface water, and 2.0 gL-1 to 10.2 gL-1 in bottom water. Cu in sediment exceeded the Effect Range Low of 34mgkg-1 (i.e. where adverse effects to fauna may occur), with a peak concentration of 442mgkg-1 at the head of the basin. Free Cu++ in surface water was several orders of magnitude higher than in sediment porewater. High-resolution data of Cu species together with probability maps presented in this paper will allow managers to easily visualise and localise areas of impaired quality and to prioritise which areas should be targeted to improve Cu-related conditions.

Levin, LA, Ziebis W, Mendoza GF, Growney VA, Tryon MD, Brown KM, Mahn C, Gieskes JM, Rathburn AE.  2003.  Spatial heterogeneity of macrofauna at northern California methane seeps: influence of sulfide concentration and fluid flow. Marine Ecology-Progress Series. 265:123-139.   10.3354/meps265123   AbstractWebsite

Relationships among fluid flow, sulfide concentration, sulfur bacteria and macrofaunal assemblages were examined at methane seeps on the northern California margin, near the mouth of the Eel River (512 to 525 m). Over a 6 mo period, sediments covered with microbial mats exhibited significant but variable outflow of altered fluids, with no flow reversals. This fluid flow was associated with high porewater sulfide concentrations (up to 20 mM) and almost no oxygen penetration of sediments (<0.1 mm). Vesicomya pacifica (clam) bed and non-seep sediments exhibited little net fluid outflow and similar oxygen penetration (3 and 4 mm, respectively); however, sulfide concentrations were higher in subsurface clam-bed sediments (up to 2 mM) than in non-seep sediments (<200 muM). Macrofaunal densities did not differ among the 3 habitats (13 800 to 16 800 ind. m(-2); >300 mum), but biomass and diversity (no. species per core, E(S-100), H') were lower and composition varied in the sulfidic microbial mat sediments relative to clam-bed and non-seep sediments. The community in microbial mat-covered sediments consisted largely (82%) of 6 species in the polychaete family Dorvilleidae, whereas the clam-bed and non-seep microhabitats supported a mixture of annelids, peracarid crustaceans, nemerteans, and mollusks. Vertical microprofiling of sulfide in animal cores indicated that most taxa avoid H2S concentrations >1 mM. However, sulfide-oxidizing filamentous bacteria, dorvilleid polychaetes and bivalves (mainly V. pacifica) exhibited highest densities at sulfide concentrations of 1 to 5 mM sulfide. Horizontal and vertical patterns of sulfide availability have a strong influence on the fine-scale distribution, structure and composition of macrofaunal assemblages inhabiting methane seeps and must be accounted for when characterizing the microbiology and ecology of seep habitats.

Zapata-Hernandez, G, Sellanes J, Thurber AR, Levin LA.  2014.  Trophic structure of the bathyal benthos at an area with evidence of methane seep activity off southern Chile (similar to 45 degrees S). Journal of the Marine Biological Association of the United Kingdom. 94:659-669.   10.1017/s0025315413001914   AbstractWebsite

Through application of carbon (C) and nitrogen (N) stable isotope analyses, we investigated the benthic trophic structure of the upper-slope off southern Chile (similar to 45 degrees S) including a recent methane seep area discovered as part of this study. The observed fauna comprised 53 invertebrates and seven fish taxa, including remains of chemosymbiotic fauna (e.g. chemosymbiotic bivalves and siboglinid polychaetes), which are typical of methane seep environments. While in close-proximity to a seep, the heterotrophic fauna had a nutrition derived predominantly from photosynthetic sources (delta C-13 > -21 parts per thousand). The absence of chemosynthesis-based nutrition in the consumers was likely a result of using an Agassiz trawl to sample the benthos, a method that is likely to collect a mix of fauna including individuals from adjacent non-seep bathyal environments. While four trophic levels were estimated for invertebrates, the fish assemblage was positioned within the third trophic level of the food web. Differences in corrected standard ellipse area (SEA(C)), which is a proxy of the isotopic niche width, yielded differences for the demersal fish Notophycis marginata (SEA(C) = 5.1 parts per thousand) and Coelorinchus fasciatus (SEA(C) = 1.1 parts per thousand), suggesting distinct trophic behaviours. No ontogenic changes were detected in C. fasciatus regarding food sources and trophic position. The present study contributes the first basic trophic data for the bathyal area off southern Chile, including the identification of a new methane seep area, among the furthest south ever discovered. Such information provides the basis for the proper sustainable management of the benthic environments present along the vast Chilean continental margin.