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2011
Ramirez-Llodra, E, Tyler PA, Baker MC, Bergstad OA, Clark MR, Escobar E, Levin LA, Menot L, Rowden AA, Smith CR, Vandover CL.  2011.  Man and the last great wilderness: human impact on the deep sea. Plos One. 6   10.1371/journal.pone.0022588   AbstractWebsite

The deep sea, the largest ecosystem on Earth and one of the least studied, harbours high biodiversity and provides a wealth of resources. Although humans have used the oceans for millennia, technological developments now allow exploitation of fisheries resources, hydrocarbons and minerals below 2000 m depth. The remoteness of the deep seafloor has promoted the disposal of residues and litter. Ocean acidification and climate change now bring a new dimension of global effects. Thus the challenges facing the deep sea are large and accelerating, providing a new imperative for the science community, industry and national and international organizations to work together to develop successful exploitation management and conservation of the deep-sea ecosystem. This paper provides scientific expert judgement and a semi-quantitative analysis of past, present and future impacts of human-related activities on global deep-sea habitats within three categories: disposal, exploitation and climate change. The analysis is the result of a Census of Marine Life - SYNDEEP workshop (September 2008). A detailed review of known impacts and their effects is provided. The analysis shows how, in recent decades, the most significant anthropogenic activities that affect the deep sea have evolved from mainly disposal (past) to exploitation (present). We predict that from now and into the future, increases in atmospheric CO(2) and facets and consequences of climate change will have the most impact on deep-sea habitats and their fauna. Synergies between different anthropogenic pressures and associated effects are discussed, indicating that most synergies are related to increased atmospheric CO(2) and climate change effects. We identify deep-sea ecosystems we believe are at higher risk from human impacts in the near future: benthic communities on sedimentary upper slopes, cold-water corals, canyon benthic communities and seamount pelagic and benthic communities. We finalise this review with a short discussion on protection and management methods.

Currin, CA, Levin LA, Talley TS, Michener R, Talley D.  2011.  The role of cyanobacteria in Southern California salt marsh food webs. Marine Ecology. 32:346-363.: Blackwell Publishing Ltd   10.1111/j.1439-0485.2011.00476.x   AbstractWebsite

Understanding wetland food webs is critical for effective habitat management, restoration and conservation. Microalgae are recognized as key food sources for marsh invertebrates but the importance of different groups under various conditions is rarely examined. We tested the hypothesis that faunal utilization of microalgae, and cyanobacteria in particular, is significant in Southern California created and natural salt marshes but varies with habitat type (creek bank versus marsh interior) and season (spring versus autumn). We used stable isotope analysis and mixing models (IsoSource) to compare food webs in adjacent young (created) and mature (natural) salt marshes. Isotopic values of some primary producers, macrofauna, epifauna, and fish demonstrated significant differences between the adjacent salt marshes. δ13C and δ34S values of the benthic microalgal community varied with taxonomic composition (diatoms versus cyanobacteria) and to a lesser extent with season. Depleted δ15N values of benthic diatoms and macroalgae indicated that N2 fixed within algal mats was recycled within the benthic algal community. Marsh fauna, including most major macrofauna taxal, Cerithidea, and Fundulus, also exhibited seasonal differences in isotopic composition, and Cerithidea and selected macrofauna (oligochaetes, polychaetes) from the marsh interior were more enriched in 13C and depleted in 15N than those from the creek bank. In the young marsh, the cyanobacteria contributed a minimum of 17–100% of the primary production in food webs supporting macrofauna, and cyanobacteria contributed at least 40% of the primary production included in Cerithidea and Fundulus food webs. A wider range of primary producers contributed to food webs in the mature marsh. Cyanobacteria were a greater source of trophic support for macrofauna from the marsh interior than the creek bank, whereas Spartina was a more important food source for creek bank macrofauna in both marshes. Insect larvae largely consumed cyanobacteria, whereas polychaetes exhibited greater utilization of Spartina. Phytoplankton was the primary food resource for mussels in both marshes. Although the spatial and temporal complexity of food webs has traditionally been collapsed into the study of relatively simplified food webs, isotope signatures reveal fine-scale patterns in food web structure that may be used to make more accurate assessments of ecosystem state. Accurate interpretation of marsh trophic structure using natural abundance stable isotopes requires fine-scale resolution in space and time, a large number of samples, and a high level of taxonomic resolution.

2010
Levin, LA, Mendoza GF, Gonzalez JP, Thurber AR, Cordes EE.  2010.  Diversity of bathyal macrofauna on the northeastern Pacific margin: the influence of methane seeps and oxygen minimum zones. Marine Ecology-an Evolutionary Perspective. 31:94-110.   10.1111/j.1439-0485.2009.00335.x   AbstractWebsite

The upper continental slope in the northeastern Pacific Ocean is intercepted by a deep oxygen minimum zone (OMZ; 650-1100 m) and punctuated by conduits of methane seepage. We examined the effects of these two dominant sources of heterogeneity on the density, composition and diversity of heterotrophic macrofauna off Hydrate Ridge, Oregon (OR; 800 m water depth), where the seeps co-occur within an OMZ, and off the Eel River, Northern California (CA; 500 m), where seeps are overlain by better oxygenated waters. We hypothesized that seeps (containing clam beds and microbial mats) should contribute a suite of distinct species to the regional margin species pool but that OMZ-associated hypoxia would dampen seep-related heterogeneity. Macrofaunal densities were highest (23,000-33,510 ind.m(-2)) in the CA seep sediments and in the OR near-seep samples, intermediate in the OR seep, CA near seep and CA and OR 500-m margin sediments (10,05419,777 ind.m(-2)), and lowest in the CA and OR OMZ habitats at 800 m (42697847 ind.m(-2)). Annelids constituted over 50% of the taxa in all but the CA clam bed and OR microbial mat sediments, where mollusks were abundant. Approximately 50% of seep species appeared to be habitat endemic; species present in microbial mats largely formed a subset of those present in the clam beds. Dorvilleid and ampharetid polychaetes were dominant in the seep sediments; non-seep margin sediments at 500 and 800 m were populated heavily by branckiate polychaetes including cossurids and paraonids. Alpha diversity (Es[20] calculated per core) was lowest and rank 1 dominance was highest in the CA and OR microbial mat habitats. Pooled analyses of Es[100] revealed highest species richness in the CA clam bed and near-seep habitats (30.3 and 29.6, respectively), and lowest species richness in the OR microbial mat and near-seep habitats (16.5 and 17.9, respectively). Non-seep sediments (500 and 800 m) off both CA and OR were more homogeneous (55-57% within-habitat similarity) than clam bed and microbial mat sediments (only 32-37% within-habitat similarity). CA sediment macrofauna generally exhibit higher alpha diversity, and as habitats are combined, a higher rate of increase in the slope of the species accumulation curves than do OR margin macrofauna. Methane seeps in the NE Pacific introduce significant heterogeneity that increases margin biodiversity at multiple spatial scales. However, our hypothesis that the OMZ would lessen the seep contributions to diversity was not supported. The better oxygenated CA seeps at 500 in shared more of the background margin fauna (at 500 m) than did the OR seeps at 800 m (with OMZ fauna at 800 in). Geographical differences in the fluxes of methane-rich fluids and the increased reliance on chemosynthetic food sources with increased depth could explain these results.

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

Ramirez-Llodra, E, Brandt A, Danovaro R, De Mol B, Escobar E, German CR, Levin LA, Arbizu PM, Menot L, Buhl-Mortensen P, Narayanaswamy BE, Smith CR, Tittensor DP, Tyler PA, Vanreusel A, Vecchione M.  2010.  Deep, diverse and definitely different: unique attributes of the world's largest ecosystem. Biogeosciences. 7:2851-2899.   10.5194/bg-7-2851-2010   AbstractWebsite

The deep sea, the largest biome on Earth, has a series of characteristics that make this environment both distinct from other marine and land ecosystems and unique for the entire planet. This review describes these patterns and processes, from geological settings to biological processes, biodiversity and biogeographical patterns. It concludes with a brief discussion of current threats from anthropogenic activities to deep-sea habitats and their fauna. Investigations of deep-sea habitats and their fauna began in the late 19th century. In the intervening years, technological developments and stimulating discoveries have promoted deep-sea research and changed our way of understanding life on the planet. Nevertheless, the deep sea is still mostly unknown and current discovery rates of both habitats and species remain high. The geological, physical and geochemical settings of the deep-sea floor and the water column form a series of different habitats with unique characteristics that support specific faunal communities. Since 1840, 28 new habitats/ecosystems have been discovered from the shelf break to the deep trenches and discoveries of new habitats are still happening in the early 21st century. However, for most of these habitats the global area covered is unknown or has been only very roughly estimated; an even smaller - indeed, minimal - proportion has actually been sampled and investigated. We currently perceive most of the deep-sea ecosystems as heterotrophic, depending ultimately on the flux on organic matter produced in the overlying surface ocean through photosynthesis. The resulting strong food limitation thus shapes deep-sea biota and communities, with exceptions only in reducing ecosystems such as inter alia hydrothermal vents or cold seeps. Here, chemoautolithotrophic bacteria play the role of primary producers fuelled by chemical energy sources rather than sunlight. Other ecosystems, such as seamounts, canyons or cold-water corals have an increased productivity through specific physical processes, such as topographic modification of currents and enhanced transport of particles and detrital matter. Because of its unique abiotic attributes, the deep sea hosts a specialized fauna. Although there are no phyla unique to deep waters, at lower taxonomic levels the composition of the fauna is distinct from that found in the upper ocean. Amongst other characteristic patterns, deep-sea species may exhibit either gigantism or dwarfism, related to the decrease in food availability with depth. Food limitation on the seafloor and water column is also reflected in the trophic structure of heterotrophic deep-sea communities, which are adapted to low energy availability. In most of these heterotrophic habitats, the dominant megafauna is composed of detritivores, while filter feeders are abundant in habitats with hard substrata (e. g. mid-ocean ridges, seamounts, canyon walls and coral reefs). Chemoautotrophy through symbiotic relationships is dominant in reducing habitats. Deep-sea biodiversity is among of the highest on the planet, mainly composed of macro and meiofauna, with high evenness. This is true for most of the continental margins and abyssal plains with hot spots of diversity such as seamounts or cold-water corals. However, in some ecosystems with particularly "extreme" physicochemical processes (e.g. hydrothermal vents), biodiversity is low but abundance and biomass are high and the communities are dominated by a few species. Two large-scale diversity patterns have been discussed for deep-sea benthic communities. First, a unimodal relationship between diversity and depth is observed, with a peak at intermediate depths (2000-3000 m), although this is not universal and particular abiotic processes can modify the trend. Secondly, a poleward trend of decreasing diversity has been discussed, but this remains controversial and studies with larger and more robust data sets are needed. Because of the paucity in our knowledge of habitat coverage and species composition, biogeographic studies are mostly based on regional data or on specific taxonomic groups. Recently, global biogeographic provinces for the pelagic and benthic deep ocean have been described, using environmental and, where data were available, taxonomic information. This classification described 30 pelagic provinces and 38 benthic provinces divided into 4 depth ranges, as well as 10 hydrothermal vent provinces. One of the major issues faced by deep-sea biodiversity and biogeographical studies is related to the high number of species new to science that are collected regularly, together with the slow description rates for these new species. Taxonomic coordination at the global scale is particularly difficult, but is essential if we are to analyse large diversity and biogeographic trends. Because of their remoteness, anthropogenic impacts on deep-sea ecosystems have not been addressed very thoroughly until recently. The depletion of biological and mineral resources on land and in shallow waters, coupled with technological developments, are promoting the increased interest in services provided by deep-water resources. Although often largely unknown, evidence for the effects of human activities in deep-water ecosystems - such as deep-sea mining, hydrocarbon exploration and exploitation, fishing, dumping and littering - is already accumulating. Because of our limited knowledge of deep-sea biodiversity and ecosystem functioning and because of the specific life-history adaptations of many deep-sea species (e. g. slow growth and delayed maturity), it is essential that the scientific community works closely with industry, conservation organisations and policy makers to develop robust and efficient conservation and management options.

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.

Schander, C, Rapp HT, Kongsrud JA, Bakken T, Berge J, Cochrane S, Oug E, Byrkjedal I, Todt C, Cedhagen T, Fosshagen A, Gebruk A, Larsen K, Levin L, Obst M, Pleijel F, Stohr S, Waren A, Mikkelsen NT, Hadler-Jacobsen S, Keuning R, Petersen KH, Thorseth IH, Pedersen RB.  2010.  The fauna of hydrothermal vents on the Mohn Ridge (North Atlantic) *. Marine Biology Research. 6:155-171.: Taylor & Francis Group Ltd., 2 Park Square Oxford OX14 4RN UK   10.1080/17451000903147450   AbstractWebsite

not available.

2009
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.

Grosholz, ED, Levin LA, Tyler C, Neira C.  2009.  Changes in community structure and ecosystem function following Spartina alterniflora invasion of Pacific estuaries. Human impacts on salt marshes : a global perspective. ( Silliman BR, Grosholz E, Bertness MD, Eds.).:23-40., Berkeley: University of California Press Abstract
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2008
Whitcraft, CR, Levin LA, Talley D, Crooks JA.  2008.  Utilization of invasive tamarisk by salt marsh consumers. Oecologia. 158:259-272.   10.1007/s00442-008-1144-5   AbstractWebsite

Plant invasions of coastal wetlands are rapidly changing the structure and function of these systems globally. Alteration of litter dynamics represents one of the fundamental impacts of an invasive plant on salt marsh ecosystems. Tamarisk species (Tamarix spp.), which extensively invade terrestrial and riparian habitats, have been demonstrated to enter food webs in these ecosystems. However, the trophic impacts of the relatively new invasion of tamarisk into marine ecosystem have not been assessed. We evaluated the trophic consequences of invasion by tamarisk for detrital food chains in the Tijuana River National Estuarine Research Reserve salt marsh using litter dynamics techniques and stable isotope enrichment experiments. The observations of a short residence time for tamarisk combined with relatively low C:N values indicate that tamarisk is a relatively available and labile food source. With an isotopic ((15)N) enrichment of tamarisk, we demonstrated that numerous macroinvertebrate taxonomic and trophic groups, both within and on the sediment, utilized (15)N derived from labeled tamarisk detritus. Infaunal invertebrate species that took up no or limited (15)N from labeled tamarisk (A. californica, enchytraeid oligochaetes, coleoptera larvae) occurred in lower abundance in the tamarisk-invaded environment. In contrast, species that utilized significant (15)N from the labeled tamarisk, such as psychodid insects, an exotic amphipod, and an oniscid isopod, either did not change or occurred in higher abundance. Our research supports the hypothesis that invasive species can alter the trophic structure of an environment through addition of detritus and can also potentially impact higher trophic levels by shifting dominance within the invertebrate community to species not widely consumed.

2007
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.

2006
Arntz, WE, Gallardo VA, Gutierrez D, Isla E, Levin LA, Mendo J, Neira C, Rowe GT, Tarazona J, Wolff M.  2006.  El Niño and similar perturbation effects on the benthos of the Humboldt, California, and Benguela Current upwelling ecosystems. Advances in Geosciences. 6:243-265.: European Geosciences Union, c/o E.O.S.T. 5, rue Rene Descartes Strasbourg Cedex 67084 France, [mailto:egu.production@copernicus.org], [URL:http://www.copernicus.org/EGU] AbstractWebsite

To a certain degree, Eastern Boundary Current (EBC) ecosystems are similar: Cold bottom water from moderate depths, rich in nutrients, is transported to the euphotic zone by a combination of trade winds, Coriolis force and Ekman transport. The resultant high primary production fuels a rich secondary production in the upper pelagic and nearshore zones, but where O sub(2) exchange is restricted, it creates oxygen minimum zones (OMZs) at shelf and upper slope (Humboldt and Benguela Current) or slope depths (California Current). These hypoxic zones host a specifically adapted, small macro- and meiofauna together with giant sulphur bacteria that use nitrate to oxydise H sub(2)S. In all EBC, small polychaetes, large nematodes and other opportunistic benthic species have adapted to the hypoxic conditions and co-exist with sulphur bacteria, which seem to be particularly dominant off Peru and Chile. However, a massive reduction of macrobenthos occurs in the core of the OMZ. In the Humboldt Current area the OMZ ranges between <100 and about 600 m, with decreasing thickness in a poleward direction. The OMZ merges into better oxygenated zones towards the deep sea, where large cold-water mega- and macrofauna occupy a dominant role as in the nearshore strip. The Benguela Current OMZ has a similar upper limit but remains shallower. It also hosts giant sulphur bacteria but little is known about the benthic fauna. However, sulphur eruptions and intense hypoxia might preclude the coexistence of significant mega- und macrobenthos. Conversely, off North America the upper limit of the OMZ is considerably deeper (e.g., 500-600 m off California and Oregon), and the lower boundary may exceed 1000m. The properties described are valid for very cold and cold (La Nina and "normal") ENSO conditions with effective upwelling of nutrient-rich bottom water. During warm (El Nino) episodes, warm water masses of low oxygen concentration from oceanic and equatorial regions enter the upwelling zones, bringing a variety of (sub)tropical immigrants. The autochthonous benthic fauna emigrates to deeper water or poleward, or suffers mortality. However, some local macrofaunal species experience important population proliferations, presumably due to improved oxygenation (in the southern hemisphere), higher temperature tolerance, reduced competition or the capability to use different food. Both these negative and positive effects of el Nino influence local artisanal fisheries and the livelihood of coastal populations. In the Humboldt Current system the hypoxic seafloor at outer shelf depths receives important flushing from the equatorial zone, causing havoc on the sulphur bacteria mats and immediate recolonisation of the sediments by mega- and macrofauna. Conversely, off California, the intruding equatorial water masses appear to have lower oxygen than ambient waters, and may cause oxygen deficiency at upper slope depths. Effects of this change have not been studied in detail, although shrimp and other taxa appear to alter their distribution on the continental margin. Other properties and reactions of the two Pacific EBC benthic ecosystems to el Nino seem to differ, too, as does the overall impact of major episodes (e.g., 1982/1983(1984) vs. 1997/1998). The relation of the "Benguela Nino" to ENSO seems unclear although many Pacific- Atlantic ocean and atmosphere teleconnections have been described. Warm, low- oxygen equatorial water seems to be transported into the upwelling area by similar mechanisms as in the Pacific, but most major impacts on the eukaryotic biota obviously come from other, independent perturbations such as an extreme eutrophication of the sediments ensuing in sulphidic eruptions and toxic algal blooms. Similarities and differences of the Humboldt and California Current benthic ecosystems are discussed with particular reference to ENSO impacts since 1972/73. Where there are data available, the authors include the Benguela Current ecosystem as another important, non-Pacific EBC, which also suffers from the effects of hypoxia.

2003
Levin, LA, Gooday A.  2003.  The Deep Atlantic Ocean. Ecosystems of the deep oceans. ( Tyler PA, Ed.).:111-178., Amsterdam ; New York: Elsevier Abstract
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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.

2002
Levin, LA, Talley TS.  2002.  Natural and manipulated sources of heterogeneity controlling early faunal development of a salt marsh. Ecological Applications. 12:1785-1802.   10.2307/3099938   AbstractWebsite

Ecosystem recovery following wetland restoration offers exceptional opportunities to study system structure, function, and successional processes in salt marshes. This study used observations of natural variation and large-scale manipulative experiments to test the influence of vascular vegetation and soil organic matter on the rate and trajectory of macrofaunal recovery in a southern California created salt marsh, the Crown Point Mitigation Site. During the first three years following marsh establishment, macrofaunal density and species richness recovered rapidly within the Spartina foliosa (cordgrass) zone; densities in the created marsh were 50% of those in the natural marsh after 16 mo and 97% after 28 mo. However, the early successional assemblage had a lower proportion of tubificid and enchytraeid oligochaetes, and a higher proportion of chironomids and other insect larvae than did the mature natural marsh. Most of the colonizers arrived by rafting on sea grass and algae rather than by larval dispersal. Initial planting of S. foliosa had no influence on macrofaunal recovery, perhaps because of variable transplant survival. However, subsequently, both positive and negative correlations were observed between densities of some macrofaunal taxa and shoot densities of S. foliosa or Salicornia spp. (pickleweed). Salinity and measures of soil organics (belowground biomass, combustible organic matter, and chlorophyll a) also were correlated with macrofaunal densities and taxon richness. Of foul added soil amendments (kelp, alfalfa, peat, and Milorganite), Milorganite (a sewage product) and kelp both promoted macrofaunal colonization during year 1, but effects were short lived. The most significant sources of heterogeneity in the recovering marsh were associated with site history and climate variation. Faunal recovery was most rapid in highly localized, organic-rich marsh sediments that were remnants of the historical wetland. Elevated sea level during the 1998 El Nino corresponded with similarity of macrofaunal communities in the created and natural marshes. The large spatial scale and multi-year duration of this study revealed that natural sources of spatial and temporal heterogeneity may exert stronger influence on faunal succession in California wetlands than manipulation of vegetation or soil properties.

Demaster, DJ, Thomas CJ, Blair NE, Fornes WL, Plaia G, Levin LA.  2002.  Deposition of bomb (14)C in continental slope sediments of the Mid-Atlantic Bight: assessing organic matter sources and burial rates. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 49:4667-4685.   10.1016/s0967-0645(02)00134-0   AbstractWebsite

As part of the Ocean Margins Program (OMP), organic carbon (14)C measurements have been made on benthic fauna and kasten core sediments from the North Carolina continental slope. These analyses are used to evaluate the nature and burial flux of organic matter in the OMP study area off Cape Hatteras. Despite the fact that surface sediment (14)C contents ranged from -41 to -215 per mil, the benthic fauna (primarily polychactes) all contained significant amounts of bomb-(14)C (body tissue (14)C contents ranging from + 20 to + 82 per mil). Bomb-(14)C clearly is reaching the seabed on the North Carolina slope, and the labile planktonic material carrying this signal is a primary source of nutrition to the benthic ecosystem. The enrichment of (14)C in benthic faunal tissue relative to the (14)C content of bulk surface-sediment organic matter (a difference of similar to 150 per mil) is attributed to a combination of particle selection and selective digestive processes. Organic carbon burial rates from 12 stations on the North Carolina slope varied from 0.02 to 1.7 mol of Cm(-2) yr(-1), with a mean value of 0.7 mol of C m(-2) yr(-1). The accumulation of organic matter on the upper slope accounts for < 1 % of the primary production in the entire continental margin system. The North Carolina margin was deliberately selected because of its potential for offshore transport and high sediment deposition rates, and even in this environment, burial of organic carbon accounts for a very small fraction of the primary production occurring in surface waters. (C) 2002 Elsevier Science Ltd. All rights reserved.

2001
Talley, TS, Crooks JA, Levin LA.  2001.  Habitat utilization and alteration by the invasive burrowing isopod, Sphaeroma quoyanum, in California salt marshes. Marine Biology. 138:561-573.   10.1007/s002270000472   AbstractWebsite

In recent years the pace of exotic species introduction and invasion has accelerated, particularly in estuaries and wetlands. Species invasions may affect coastal ecosystems in many ways. Alteration of sedimentary environments, through structure formation and burrowing, has particularly dramatic effects on coastal habitats. This study examines modification of channel bank and marsh edge habitat by the burrowing Australasian isopod Sphaeroma quoyanum Milne Edwards, in created and natural salt marshes of San Diego Bay and San Francisco Bay. Abundance and distribution patterns of this isopod species, its relationships with habitat characteristics, and its effects on sediment properties and bank erosion were examined seasonally, and in several marsh microhabitats. Mean isopod densities were 1541 and 2936 individuals per 0.25 m(2) in San Francisco Bay, and 361 and 1153 individuals per 0.25 m(2) in San Diego Bay study sites during December and July 1998, respectively. This isopod forms dense, anastomosing burrow networks. S. quoyanum densities did not differ as a function of location within creeks or location in natural versus created marshes. Burrows, which are on average 6 mm wide and 2 cm long, were associated with firm sediments containing high detrital biomass. Although erosion is a natural process along salt marsh banks, enclosure experiments demonstrated that isopod activities can enhance sediment loss from banks. In areas infested with S. quoyanum, losses may exceed 100 cm of marsh edge per year. The effects of habitat alteration by this invading species are likely to increase in severity in the coastal zone as these ecosystems become degraded.

Blair, NE, Levin LA, Demaster DJ, Plaia G, Martin C, Fornes W, Thomas C, Pope R.  2001.  The biogeochemistry of carbon in continental slope sediments. Organism-sediment Interactions. ( Aller JY, Woodin S, Aller RC, Belle W. Baruch Institute for Marine Biology and Coastal Research. , Eds.).:243-262., Columbia: Published for the Belle W. Baruch Insitute for Marine Biology and Coastal Research by the University of South Carolina Press Abstract
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Talley, TS, Levin LA.  2001.  Modification of sediments and macrofauna by an invasive marsh plant. Biological Invasions. 3:51-68.   10.1023/A:1011453003168   AbstractWebsite

Invasive grasses have recently altered salt marsh ecosystems throughout the northern hemisphere. On the eastern seaboard of the USA, Phragmites australis has invaded both brackish and salt marsh habitats. Phragmites australis influence on sediments and fauna was investigated along a salinity and invasion-age gradient in marshes of the lower Connecticut River estuary. Typical salinities were about 19-4 ppt in Site I, 9-10 ppt in Site II and 5-7 ppt in Site III. Strongest effects were evident in the least saline settings (II and III) where Phragmites has been present the longest and exists in monoculture. Limited influence was evident in the most saline region (I) where Phragmites and native salt marsh plants co-occur. The vegetation within Phragmites stands in tidal regions of the Connecticut River generally exhibits taller, but less dense shoots, higher above-ground biomass, and lower below-ground biomass than does the un-invaded marsh flora. There were lower sediment organic content, greater litter accumulation and higher sediment chlorophyll a concentrations in Phragmites-invaded than un-invaded marsh habitat. Epifaunal gastropods (Succinea wilsoni and Stagnicola catascopium) were less abundant in habitats where Phragmites had invaded than in un-invaded marsh habitat. Macro-infaunal densities were lower in the Phragmites-invaded than un-invaded habitats at the two least saline sites (II and III). Phragmites stands supported more podurid insects, sabellid polychaetes, and peracarid crustaceans, fewer arachnids, midges, tubificid and enchytraeid oligochaetes, and greater habitat-wide taxon richness as measured by rarefaction, than did the uninvaded stands. The magnitude and significance of the compositional differences varied with season and with site; differences were generally greatest at the oldest, least saline sites (II and III) and during May, when faunal densities were higher than in September. However, experimental design and the 1-year study period precluded clear separation of salinity, age, and seasonal effects. Although structural effects of Phragmites on salt marsh faunas are evident, further investigation is required to determine the consequences of these effects for ecosystem function.

2000
Levin, LA, Talley T.  2000.  Influences of vegetation and abiotic environmental factors on salt marsh benthos. Concepts and controversies in tidal marsh ecology. ( Weinstein M, Kreeger D, Eds.).:661-708., Dordrecht ; Boston: Kluwer Academic Abstract
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1999
Talley, TS, Levin LA.  1999.  Macrofaunal succession and community structure in Salicornia marshes of southern California. Estuarine Coastal and Shelf Science. 49:713-731.   10.1006/ecss.1999.0553   AbstractWebsite

Lack of basic understanding of ecosystem structure and function forms a major impediment to successful conservation of coastal ecosystems. This paper provides a description of the fauna and examines faunal succession in Salicornia-vegetated sediments of southern California. Environmental attributes (vegetation and sediment properties) and macrofaunal (animals greater than or equal to 0.3 mm) community structure were examined in sediments of five natural, southern California Salicornia spp. marshes (Tijuana Estuary, San Diego Bay, Mission Bay, Upper Newport Bay and Anaheim Bay) and in created Salicornia marshes 16 months to 10 years in age, located within four of the bays. Oligochaetes and insects were the dominant taxa in both natural (71 to 98% of total fauna) and created (91 to 97%) marshes. In San Diego, Newport and Anaheim Bays, macrofaunal densities were generally higher in the created marshes (88 000 to 290 000 ind m(-2)) than in their natural counterparts (26 000 to 50 000 ind m(-2)). In the youngest system, Mission Bay, the reverse was true (natural: 113 000 vs created: 28 000 ind m-2). Similar species numbers were recorded from the created and adjacent natural marshes. Insects, especially chironomids, dolichopodids, and heleids, as well as the naidid oligochaete, Paranais litoralis, characterize early successional stages. Enchytraeid and tubificid oligochaetes reflect later succession evident in natural and older created marshes. Sediment organic matter (both combustible and below-ground plant biomass) was the environmental variable most commonly associated with densities of various macrofaunal taxa. These relationships were generally negative in the natural marshes and positive in the created marshes. Within-bay comparisons of macrofauna from natural Salicornia- vs Spartina-vegetated habitat in San Diego and Mission Bays revealed lower macrofaunal density (San Diego Bay only), proportionally fewer oligochaetes and more insects, and no differences in species richness in the Salicornia habitat. The oldest created Salicornia marsh (San Diego Bay) exhibited an assemblage intermediate in composition between those of the natural Salicornia- and Spartina-vegetated marshes. These results suggest: (a) faunal recovery following Salicornia marsh creation can require 10 or more years, (b) high macrofaunal variability among bays requires marsh creation reference site selection from within the same bay, and (c) Spartina-based research should not be used for Salicornia marsh management decisions. (C) 1999 Academic Press.

Thistle, D, Levin LA, Gooday AJ, Pfannkuche O, Lambshead PJD.  1999.  Physical reworking by near-bottom flow alters the metazoan meiofauna of Fieberling Guyot (northeast Pacific). Deep-Sea Research Part I-Oceanographic Research Papers. 46:2041-2052.   10.1016/s0967-0637(99)00040-0   AbstractWebsite

Although much of the deep sea is physically tranquil, some regions experience near-bottom flows that rework the surficial sediment. During periods of physical reworking, animals in the reworked layer risk being suspended, which can have both positive and negative effects. Reworking can also change the sediment in ecologically important ways, so the fauna of reworked sites should differ from that of quiescent locations. We combined data from two reworked, bathyal sites on the summit of Fieberling Guyot (32 degrees 27.631'N, 127 degrees 49.489'W; 32 degrees 27.581'N, 127 degrees 47.839'W) and compared the results with those of more tranquil sites. We tested for differences in the following parameters, which seemed likely to be sensitive to the direct or indirect effects of reworking: (1) the vertical distribution of the meiofauna in the sea bed, (2) the relative abundance of surface-living harpacticoids, (3) the proportion of the fauna consisting of interstitial harpacticoids, (4) the ratio of harpacticoids to nematodes. We found that the vertical distributions of harpacticoid copepods, ostracods, and kinorhynchs were deeper on Fieberling. In addition, the relative abundance of surface-living harpacticoids was less, the proportion of interstitial harpacticoids was greater, and the ratio of harpacticoids to nematodes was greater on Fieberling. These differences between Fieberling and the comparison sites suggest that physical reworking affects deep-sea meiofauna and indicate the nature of some of the effects. (C) 1999 Elsevier Science Ltd. AII rights reserved.

Levin, LA, Blair NE, Martin CM, Demaster DJ, Plaia G, Thomas CJ.  1999.  Macrofaunal processing of phytodetritus at two sites on the Carolina margin: in situ experiments using (13)C-labeled diatoms. Marine Ecology-Progress Series. 182:37-54.   10.3354/meps182037   AbstractWebsite

Tracer experiments using (13)C-labeled diatoms Thalassiosira pseudonana were carried out at two 850 m sites (I off Cape Fear and III off Cape Hatteras) on the North Carolina, USA, slope to examine patterns of macrofaunal consumption of fresh phytodetritus. Experiments examined the influence of taxon, feeding mode, body size and vertical position within the sediment column on access to surficial organic matter. delta(13)C measurements were made on macrofaunal metazoans and agglutinating protozoans from background sediments and from sediment plots in which (13)C-labeled diatoms were deposited and then sampled 0.3 h, 1 to 1.5 d, 3 mo and 14 mo later. Significant between-site differences were observed in background delta(13)C signatures of sediments, metazoans, and large, agglutinating protozoans, with values 2 to 3 parts per thousand lower at Site III than at Site I. Background delta(13)C signatures also varied as a function of taxon and of vertical position in the sediment column at Site III. The background delta(13)C value of carnivores was higher than that of surface-deposit feeders among Site I annelids, but no annelid feeding-group differences were observed at Site III. delta(13)C data from short-term (1 to 1.5 d) experiments revealed rapid diatom ingestion, primarily by agglutinated protozoans and annelids at Site I and mainly by annelids at Site III. Selective feeding on diatoms was exhibited by paraonid polychaetes, especially Aricidea spp. Exceptionally high uptake and retention of diatom C also was observed in the maldanid Praxillella sp., the nereid Ceratocephale sp. and several other surface-deposit feeding polychaetes. After 14 mo, little of the diatom (13)C remained at Site III, but high concentrations of the tracer were present in annelids and agglutinating protozoans at Site I. At both sites, nonannelid metazoans and subsurface-deposit feeding annelids exhibited the least uptake and retention of diatom C. Our hypotheses that large-bodied taxa and shallow-dwelling infauna should have greatest access to freshly deposited organic matter were not borne out. Some small, deep-dwelling taxa acquired label more readily than large or near-surface forms. Differences in tracer fates between sites reflected greater vertical mixing at Site III. These results indicate heterogeneity in benthic processes along the Carolina margin. but suggest that labile organic matter is consumed quickly at both sites. Because most of the taxa found to consume freshly deposited diatoms in these experiments are typical of bathyal settings, we infer that phytodetritus reaching the seabed in margin environments is rapidly processed by protozoan and metazoan components of the benthic fauna.

1998
Levin, LA, Talley TS, Hewitt J.  1998.  Macrobenthos of Spartina foliosa (Pacific cordgrass) salt marshes in southern California: Community structure and comparison to a Pacific mudflat and a Spartina alterniflora (Atlantic smooth cordgrass) marsh. Estuaries. 21:129-144.   10.2307/1352552   AbstractWebsite

Environmental attributes (vegetation and sediment properties) of and macrofaunal community structure in sediments of five southern California Spartina foliosa marshes (San Diego Bay, Mission Bay, Upper Newport Bay, Bolsa Chica Lagoon, and Anaheim Bay) were examined during October 1994. Macrofaunal densities in Pacific S. foliosa marshes (avg. 122,268 indiv. m(-2) > 300 mu m) were 3 to 10 times higher than observed in Atlantic S. alterniflora and S. anglica marshes. The macrofauna of S. foliosa marshes was composed mainly of enchytraeid, naidid, and tubificid oligochaetes (66%), with the enchytraeids dominant at all sites except Bolsa Chica Lagoon. Polychaetes, insects, and peracarid crustaceans accounted for most of the remaining fauna. Multivariate analyses indicated greatest faunal similarity between the two southernmost marshes (Mission Bay and San Diego Bay), and between Anaheim and Newport Bay marshes, with Bolsa Chica Lagoon exhibiting a distinct assemblage. There were strong positive associations of faunal abundance and composition with percent organic matter and percent open area, and negative associations with percent sand and dry weight of algae. For the vegetated marsh in Mission Bay, faunal comparisons were made with an adjacent mudflat and with a S. alterniflora marsh in North Carolina, USA. The unvegetated mudflat exhibited similar macrofaunal densities but higher species richness than the adjacent Spartina marsh. The macrofaunal assemblage of the Mission Bay S. foliosa marsh differed from that of the Atlantic S. alterniflora marsh and the Pacific mudflat in having a greater proportion of oligochaetes, especially Enchytraeidae, and fewer polychaetes. This study represents the first published description that we are aware of for macrofauna in S. foliosa vegetated marsh sediments. The findings document faunal variation among southern California embayments and suggest that differences in macrobenthic community structure occur between marsh and mudflat habitat as web as between east and west coast Spartina marshes. Observed differences may have significant implications for wetland conservation and restoration efforts.

Thistle, D, Levin LA.  1998.  The effect of experimentally increased near-bottom flow on metazoan meiofauna at a deep-sea site, with comparison data on macrofauna. Deep-Sea Research Part I-Oceanographic Research Papers. 45:625-+.   10.1016/s0967-0637(97)00101-5   AbstractWebsite

It has been argued that strong near-bottom hows affect macrofauna and meiofauna in the deep sea, but the evidence comes largely from studies that compared sites separated geographically by hundreds to thousands of kilometers and in depth by hundreds of meters. In this paper, the results of the first experimental investigation of the effects of strong near-bottom flow on deep-sea metazoan meiofauna are presented. At a site (32 degrees 27.581' N, 127 degrees 47.839' W) at 583 m depth on the Fieberling Guyot summit plain, the submersible Alvin emplaced weirs designed to increase the near-bottom flow locally. After 6.5 weeks, sediments in the weirs and unmanipulated locations in the vicinity were sampled. The abundances of nematodes, harpacticoid copepods, ostracods, and kinorhynchs, considered collectively and as individual taxa, were significantly lower in the weir samples than in the background samples. Parallel responses were observed in total macrofaunal and mollusk abundances. Proportional declines in kinorhynchs and mollusks were observed as well. These results suggest that strong near-bottom flow can reduce the abundance of meiofauna and macrofauna in the deep sea and alter assemblage composition. (C) 1998 Elsevier Science Ltd. All rights reserved.