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Book Chapter
Springer, AE, Stevens LE, Anderson DE, Partnell RA, Kreamer DK, Levin LA, Flora S.  2008.  A comprehensive springs classification system. Integrating geomorphic, hydrogeochemical, and ecological criteria. Aridland springs in North America: ecology and conservation. ( Stevens LE, Meretsky VJ, Eds.).:49-75., Tucson: University of Arizona Press and the Arizona-Sonora Desert Museum Abstract
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
Levin, LA, Nittrouer CA.  1987.  Textural characteristics of sediments on deep seamounts in the eastern Pacific Ocean between 10°N and 30°N. Seamounts, islands, and atolls. ( Keating BH, Fryer P, Batiza R, Boehlert GW, Eds.).:187-203., Washington, D.C.: American Geophysical Union Abstract
Journal Article
Breitburg, DL, Salisbury J, Bernhard JM, Cai WJ, Dupont S, Doney SC, Kroeker KJ, Levin LA, Long WC, Milke LM, Miller SH, Phelan B, Passow U, Seibel BA, Todgham AE, Tarrant AM.  2015.  And on top of all that... Coping with ocean acidification in the midst of many stressors. Oceanography. 28:48-61.   10.5670/oceanog.2015.31   AbstractWebsite

Oceanic and coastal waters are acidifying due to processes dominated in the open ocean by increasing atmospheric CO2 and dominated in estuaries and some coastal waters by nutrient-fueled respiration. The patterns and severity of acidification, as well as its effects, are modified by the host of stressors related to human activities that also influence these habitats. Temperature, deoxygenation, and changes in food webs are particularly important co-stressors because they are pervasive, and both their causes and effects are often mechanistically linked to acidification. Development of a theoretical underpinning to multiple stressor research that considers physiological, ecological, and evolutionary perspectives is needed because testing all combinations of stressors and stressor intensities experimentally is impossible. Nevertheless, use of a wide variety of research approaches is a logical and promising strategy for improving understanding of acidification and its effects. Future research that focuses on spatial and temporal patterns of stressor interactions and on identifying mechanisms by which multiple stressors affect individuals, populations, and ecosystems is critical. It is also necessary to incorporate consideration of multiple stressors into management, mitigation, and adaptation to acidification and to increase public and policy recognition of the importance of addressing acidification in the context of the suite of other stressors with which it potentially interacts.

Buhl-Mortensen, L, Vanreusel A, Gooday AJ, Levin LA, Priede IG, Buhl-Mortensen P, Gheerardyn H, King NJ, Raes M.  2010.  Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. Marine Ecology-an Evolutionary Perspective. 31:21-50.   10.1111/j.1439-0485.2010.00359.x   AbstractWebsite

Biological structures exert a major influence on species diversity at both local and regional scales on deep continental margins. Some organisms use other species as substrates for attachment, shelter, feeding or parasitism, but there may also be Mutual benefits from the association. Here, we highlight the structural attributes and biotic effects of the habitats that corals, sea pens, sponges and xenophyophores offer other organisms. The environmental setting of the biological structures influences their species composition. The importance of benthic species as substrates seems to increase with depth as the complexity of the surrounding geological substrate and food supply decline. There are marked differences in the degree of mutualistic relationships between habitat-forming taxa. This is especially evident for scleractinian corals, which have high numbers of facultative associates (commensals) and few obligate associates (mutualists), and gorgonians, with their few commensals and many obligate associates. Size, flexibility and architectural complexity of the habitat-forming organism are positively related to species diversity for both sessile and mobile species. This is mainly evident for commensal species sharing a facultative relationship with their host. Habitat complexity is enhanced by the architecture of biological structures, as well as by biological interactions. Colony morphology has a great influence on feeding efficiency for suspension feeders. Suspension feeding, habitat-forming organisms modify the environment to optimize their food uptake. This environmental advantage is also passed on to associated filter-feeding species. These effects are poorly understood but represent key points for understanding ecosystems and biodiversity on continental margins. In this paper we explore the contributions of organisms and the biotic structures they create (rather than physical modifications) to habitat heterogeneity and diversity on the deep continental margins.

Mengerink, KJ, Vandover CL, Ardron J, Baker M, Escobar-Briones E, Gjerde K, Koslow J, Ramirez-Llodra E, Lara-Lopez A, Squires D, Sutton T, Sweetman A, Levin LA.  2014.  A Call For Deep-Ocean Stewardship. Science. 344:696-698.
Basak, C, Rathburn AE, Perez ME, Martin JB, Kluesner JW, Levin LA, De Deckker P, Gieskes JM, Abriani M.  2009.  Carbon and oxygen isotope geochemistry of live (stained) benthic foraminifera from the Aleutian Margin and the Southern Australian Margin. Marine Micropaleontology. 70:89-101.   10.1016/j.marmicro.2008.11.002   AbstractWebsite

Comparisons of ambient bottom-water geochemistry and stable isotopic values of the tests of living (stained) calcareous benthic foraminifera from the North Pacific (on the Aleutian Margin, water depth 1988 m) and Murray Canyons group in the Southern Indian Ocean (Australian Margin, water depths 2476 m and 1634 m) provide modem environmental analogs to calibrate paleoenvironmental assessments. Consistent with the hypothesis that microhabitat preferences influence foraminiferal isotopic values, benthic foraminifera from both margins were depleted in (13)C with respect to bottom-water dissolved inorganic carbon (DIC). The carbon isotope values of deep infaunal foraminifera (Chilostomella oolina, Globobulimina pacifica) showed greater differences from estimates of those of DIC than shallow benthic foraminifera (Bulimina mexicana, Bolivinita quadrilatera, Pullenia bulloides). This study provides new isotopic and ecological information for B. quadrilatera. The mean Delta delta(13)C value, defined as foraminiferal delta(13)C values minus estimated ambient delta(13)C values from the Aleutian Margin, is 0.97 parts per thousand higher for G. pacifica than the mean from the Murray Canyon. This difference may result either from genetic or biological differences between the populations or from differences in environmental isotopic influences (such as pore water differences) that were not accounted for in the equilibrium calculations. These analyses provide calibration information for the evaluation of bottom water conditions and circulation patterns of ancient oceans based on fossil foraminiferal geochemistry. (C) 2008 Elsevier B.V. All rights reserved.

Navarro, MO, Kwan GT, Batalov O, Choi CY, Pierce NT, Levin LA.  2016.  Development of embryonic market squid, Doryteuthis opalescens, under chronic exposure to low environmental pH and O-2. Plos One. 11   10.1371/journal.pone.0167461   AbstractWebsite

The market squid, Doryteuthis opalescens, is an important forage species for the inshore ecosystems of the California Current System. Due to increased upwelling and expansion of the oxygen minimum zone in the California Current Ecosystem, the inshore environment is expected to experience lower pH and [O-2] conditions in the future, potentially impacting the development of seafloor-attached encapsulated embryos. To understand the consequences of this co-occurring environmental pH and [O-2] stress for D. opalescens encapsulated embryos, we performed two laboratory experiments. In Experiment 1, embryo capsules were chronically exposed to a treatment of higher (normal) pH (7.93) and [O-2] (242 mu M) or a treatment of low pH (7.57) and [O-2] (80 mu M), characteristic of upwelling events and/or La Nina conditions. The low pH and low [O-2] treatment extended embryo development duration by 5-7 days; embryos remained at less developed stages more often and had 54.7% smaller statolith area at a given embryo size. Importantly, the embryos that did develop to mature embryonic stages grew to sizes that were similar (non-distinct) to those exposed to the high pH and high [O-2] treatment. In Experiment 2, we exposed encapsulated embryos to a single stressor, low pH (7.56) or low [O-2] (85 mu M), to understand the importance of environmental pH and [O-2] rising and falling together for squid embryogenesis. Embryos in the low pH only treatment had smaller yolk reserves and bigger statoliths compared to those in low [O-2] only treatment. These results suggest that D. opalescens developmental duration and statolith size are impacted by exposure to environmental [O-2] and pH (pCO(2)) and provide insight into embryo resilience to these effects.

Mullineaux, LS, Metaxas A, Beaulieu SE, Bright M, Gollner S, Grupe BM, Herrera S, Kellner JB, Levin LA, Mitarai S, Neubert MG, Thurnherr AM, Tunnicliffe V, Watanabe HK, Won YJ.  2018.  Exploring the ecology of deep-sea hydrothermal vents in a metacommunity framework. Frontiers in Marine Science. 5   10.3389/fmars.2018.00049   AbstractWebsite

Species inhabiting deep-sea hydrothermal vents are strongly influenced by the geological setting, as it provides the chemical-rich fluids supporting the food web, creates the patchwork of seafloor habitat, and generates catastrophic disturbances that can eradicate entire communities. The patches of vent habitat host a network of communities (a metacommunity) connected by dispersal of planktonic larvae. The dynamics of the metacommunity are influenced not only by birth rates, death rates and interactions of populations at the local site, but also by regional influences on dispersal from different sites. The connections to other communities provide a mechanism for dynamics at a local site to affect features of the regional biota. In this paper, we explore the challenges and potential benefits of applying metacommunity theory to vent communities, with a particular focus on effects of disturbance. We synthesize field observations to inform models and identify data gaps that need to be addressed to answer key questions including: (1) what is the influence of the magnitude and rate of disturbance on ecological attributes, such as time to extinction or resilience in ametacommunity; (2) what interactions between local and regional processes control species diversity, and (3) which communities are "hot spots" of key ecological significance. We conclude by assessing our ability to evaluate resilience of vent metacommunities to human disturbance (e.g., deep-sea mining). Although the resilience of a few highly disturbed vent systems in the eastern Pacific has been quantified, these values cannot be generalized to remote locales in the western Pacific ormid Atlantic where disturbance rates are different and information on local controls is missing.

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.

Levin, LA, Boesch DF, Covich A, Dahm C, Erseus C, Ewel KC, Kneib RT, Moldenke A, Palmer MA, Snelgrove P, Strayer D, Weslawski JM.  2001.  The function of marine critical transition zones and the importance of sediment biodiversity. Ecosystems. 4:430-451.   10.1007/s10021-001-0021-4   AbstractWebsite

Estuaries and coastal wetlands are critical transition zones (CTZs) that link land, freshwater habitats, and the sea. CTZs provide essential ecological functions, including decomposition, nutrient cycling, and nutrient production, as well as regulation of fluxes of nutrients, water, particles, and organisms to and from land, rivers, and the ocean. Sediment-associated biota are integral to these functions. Functional groups considered essential to CTZ processes include heterotrophic bacteria and fungi, as well as many benthic invertebrates. Key invertebrate functions include shredding, which breaks down and recycles organic matter; suspension feeding, which collects and transports sediments across the sediment-water interface; and bioturbating, which moves sediment into or out of the seabed. In addition, macrophytes regulate many aspects of nutrient, particle, and organism dynamics above- and belowground. Animals moving within or through CTZs are vectors that transport nutrients and organic matter across terrestrial, freshwater, and marine interfaces. Significant threats to biodiversity within CTZs are posed by anthropogenic influences; eutrophication, nonnutrient pollutants, species invasions, overfishing, habitat alteration, and climate change affect species richness or composition in many coastal environments. Because biotic diversity in marine CTZ sediments is inherently low whereas their functional significance is great, shifts in diversity are likely to be particularly important. Species introductions (from invasion) or loss (from overfishing or habitat alteration) provide evidence that single-species changes can have overt, sweeping effects on CTZ structure and function. Certain species may be critically important to the maintenance of ecosystem functions in CTZs even though at present there is limited empirical evidence that the number of species in CTZ sediments is critical. We hypothesized that diversity is indeed important to ecosystem function in marine CTZs because high diversity maintains positive interactions among species (facilitation and mutualism), promoting stability and resistance to invasion or other forms of disturbance. The complexity of interactions among species and feedbacks with ecosystem functions suggests that comparative (mensurative) and manipulative approaches will be required to elucidate the role of diversity in sustaining CTZ functions.

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

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

Levin, LA, Bett BJ, Gates AR, Heimbach P, Howe BM, Janssen F, McCurdy A, Ruhl HA, Snelgrove P, Stocks KI, Bailey D, Baumann-Pickering S, Beaverson C, Benfield MC, Booth DJ, Carreiro-Silva M, Colaco A, Eble MC, Fowler AM, Gjerde KM, Jones DOB, Katsumata K, Kelley D, Le Bris N, Leonardi AP, Lejzerowicz F, Macreadie PI, McLean D, Meitz F, Morato T, Netburn A, Pawlowski J, Smith CR, Sun S, Uchida H, Vardaro MF, Venkatesan R, Weller RA.  2019.  Global observing needs in the deep ocean. Frontiers in Marine Science. 6   10.3389/fmars.2019.00241   AbstractWebsite

The deep ocean below 200 m water depth is the least observed, but largest habitat on our planet by volume and area. Over 150 years of exploration has revealed that this dynamic system provides critical climate regulation, houses a wealth of energy, mineral, and biological resources, and represents a vast repository of biological diversity. A long history of deep-ocean exploration and observation led to the initial concept for the Deep-Ocean Observing Strategy (DOOS), under the auspices of the Global Ocean Observing System (GOOS). Here we discuss the scientific need for globally integrated deep-ocean observing, its status, and the key scientific questions and societal mandates driving observing requirements over the next decade. We consider the Essential Ocean Variables (EOVs) needed to address deep-ocean challenges within the physical, biogeochemical, and biological/ecosystem sciences according to the Framework for Ocean Observing (FOO), and map these onto scientific questions. Opportunities for new and expanded synergies among deep-ocean stakeholders are discussed, including academic-industry partnerships with the oil and gas, mining, cable and fishing industries, the ocean exploration and mapping community, and biodiversity conservation initiatives. Future deep-ocean observing will benefit from the greater integration across traditional disciplines and sectors, achieved through demonstration projects and facilitated reuse and repurposing of existing deep-sea data efforts. We highlight examples of existing and emerging deep-sea methods and technologies, noting key challenges associated with data volume, preservation, standardization, and accessibility. Emerging technologies relevant to deep-ocean sustainability and the blue economy include novel genomics approaches, imaging technologies, and ultra-deep hydrographic measurements. Capacity building will be necessary to integrate capabilities into programs and projects at a global scale. Progress can be facilitated by Open Science and Findable, Accessible, Interoperable, Reusable (FAIR) data principles and converge on agreed to data standards, practices, vocabularies, and registries. We envision expansion of the deep-ocean observing community to embrace the participation of academia, industry, NGOs, national governments, international governmental organizations, and the public at large in order to unlock critical knowledge contained in the deep ocean over coming decades, and to realize the mutual benefits of thoughtful deep-ocean observing for all elements of a sustainable ocean.

Hofmann, GE, Smith JE, Johnson KS, Send U, Levin LA, Micheli F, Paytan A, Price NN, Peterson B, Takeshita Y, Matson PG, Crook ED, Kroeker KJ, Gambi MC, Rivest EB, Frieder CA, Yu PC, Martz TR.  2011.  High-Frequency Dynamics of Ocean pH: A Multi-Ecosystem Comparison. Plos One. 6   10.1371/journal.pone.0028983   AbstractWebsite

The effect of Ocean Acidification (OA) on marine biota is quasi-predictable at best. While perturbation studies, in the form of incubations under elevated pCO(2), reveal sensitivities and responses of individual species, one missing link in the OA story results from a chronic lack of pH data specific to a given species' natural habitat. Here, we present a compilation of continuous, high-resolution time series of upper ocean pH, collected using autonomous sensors, over a variety of ecosystems ranging from polar to tropical, open-ocean to coastal, kelp forest to coral reef. These observations reveal a continuum of month-long pH variability with standard deviations from 0.004 to 0.277 and ranges spanning 0.024 to 1.430 pH units. The nature of the observed variability was also highly site-dependent, with characteristic diel, semi-diurnal, and stochastic patterns of varying amplitudes. These biome-specific pH signatures disclose current levels of exposure to both high and low dissolved CO2, often demonstrating that resident organisms are already experiencing pH regimes that are not predicted until 2100. Our data provide a first step toward crystallizing the biophysical link between environmental history of pH exposure and physiological resilience of marine organisms to fluctuations in seawater CO2. Knowledge of this spatial and temporal variation in seawater chemistry allows us to improve the design of OA experiments: we can test organisms with a priori expectations of their tolerance guardrails, based on their natural range of exposure. Such hypothesis-testing will provide a deeper understanding of the effects of OA. Both intuitively simple to understand and powerfully informative, these and similar comparative time series can help guide management efforts to identify areas of marine habitat that can serve as refugia to acidification as well as areas that are particularly vulnerable to future ocean change.

Hunter, WR, Levin LA, Kitazato H, Witte U.  2012.  Macrobenthic assemblage structure and organismal stoichiometry control faunal processing of particulate organic carbon and nitrogen in oxygen minimum zone sediments. Biogeosciences. 9:993-1006.   10.5194/bg-9-993-2012   AbstractWebsite

The Arabian Sea oxygen minimum zone (OMZ) impinges on the western Indian continental margin between 150 and 1500 m, causing gradients in oxygen availability and sediment geochemistry at the sea floor. Oxygen availability and sediment geochemistry are important factors structuring macrofaunal assemblages in marine sediments. However, relationships between macrofaunal assemblage structure and sea-floor carbon and nitrogen cycling are poorly understood. We conducted in situ C-13:N-15 tracer experiments in the OMZ core (540 m [O-2] = 0.35 mu mol l(-1)) and lower OMZ boundary (800-1100 m, [O-2] = 2.2-15.0 mu mol l(-1)) to investigate how macrofaunal assemblage structure, affected by different oxygen levels, and C:N coupling influence the fate of particulate organic matter. No macrofauna were present in the OMZ core. Within the OMZ boundary, relatively high abundance and biomass resulted in the highest macrofaunal assimilation of particulate organic carbon (POC) and nitrogen (PON) at the lower oxygen 800 m stations ([O-2] = 2.2-2.36 mu mol l(-1)). At these stations the numerically dominant cirratulid polychaetes exhibited greatest POC and PON uptake. By contrast, at the higher oxygen 1100 m station ([O-2] = 15.0 mu mol l(-1)) macrofaunal C and N assimilation was lower, with POC assimilation dominated by one large solitary ascidian. Macrofaunal POC and PON assimilation were influenced by changes in oxygen availability, and significantly correlated to differences in macrofaunal assemblage structure between stations. However, macrofaunal feeding responses were ultimately characterised by preferential organic nitrogen assimilation, relative to their internal C:N budgets.

Levin, LA, Mendoza GF, Konotchick T, Lee R.  2009.  Macrobenthos community structure and trophic relationships within active and inactive Pacific hydrothermal sediments. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 56:1632-1648.   10.1016/j.dsr2.2009.05.010   AbstractWebsite

Hydrothermal fluids passing through sediments create a habitat hypothesized to influence the community structure of infaunal macrobenthos. Here we characterize the density, biomass, species composition, diversity, distributions, lifestyle, and nutritional sources of macroinfauna in hydrothermal sediments in NE and SW Pacific settings, and draw comparisons in search of faunal attributes characteristic of this habitat. There is increasing likelihood that seafloor massive sulfide deposits, associated with active and inactive hydrothermal venting, will be mined commercially. This creates a growing imperative for a more thorough understanding of the structure, dynamics, and resilience of the associated sediment faunas, and has stimulated the research presented here. Macrobenthic assemblages were studied at Manus Basin (1430-1634 m, Papua New Guinea [PNG]) as a function of location (South Su vs. Solwara 1), and hydrothermal activity (active vs. inactive), and at Middle Valley (2406-2411 m, near Juan de Fuca Ridge) as a function of habitat (active clam bed, microbial mat, hot mud, inactive background sediment). The studies conducted in PNG formed part of the environmental impact assessment work for the Solwara 1 Project of Nautilus Minerals Niugini Limited. We hypothesized that hydrothermally active sites should support (a) higher densities and biomass, (b) greater dominance and lower diversity, (c) a higher fraction of deposit feeders, and (d) greater isotopic evidence for chemosynthetic food sources than inactive sites. Manus Basin macrofauna generally had low density (<1000ind.m(-2)) and low biomass (0.1-1.07gm(-2)), except for the South Su active site, which had higher density (3494ind.m(-2)) and biomass (11.94gm(-2)), greater dominance (R1D=76%), lower diversity and more spatial (between-core) homogeneity than the Solwara 1 and South Su inactive sites. Dominant taxa at Manus Basin were Spionidae (Prionospio sp.) in active sediments, and tanaids and deposit-feeding nuculanoid bivalves in active and inactive sediments. At Middle Valley, hot mud sediments supported few animals (1011 ind m(-2)) and low biomass (1.34g m(-2)), while active clam bed sediments supported a high-density (19,984indm(-2)), high-biomass (4.46gm(-2)), low-diversity assemblage comprised of largely orbiniid and syllid polychaetes. Microbial mat sediments had the most diverse assemblage (mainly orbiniid, syllid, dorvilleid, and ampharetid polychaetes) with intermediate densities (8191 ind m(-2)) and high biomass (4.23 g m(-2)). Fauna at both Manus Basin active sites had heavy delta(13)C signatures (-17 parts per thousand to -13 parts per thousand) indicative of chemosynthetic, TCA-cycle microbes at the base of the food chain. In contrast, photosynthesis and sulfide oxidation appear to fuel most of the fauna at Manus Basin inactive sites (delta(13)C = -29 parts per thousand to -20 parts per thousand) and Middle Valley active clam beds and microbial mats (delta(13)C = -36 parts per thousand to -20 parts per thousand). The two hydrothermal regions, located at opposite ends of the Pacific Ocean, supported different habitats, sharing few taxa at the generic or family level, but both exhibited elevated infaunal density and high dominance at selected sites. Subsurface-deposit feeding and bacterivory were prevalent feeding modes. Both the Manus Basin and Middle Valley assemblages exhibit significant within-region heterogeneity, apparently conferred by variations in hydrothermal activity and associaed biogenic habitats. (C) 2009 Elsevier Ltd. All rights reserved.

Levin, LA, McGregor AL, Mendoza GF, Woulds C, Cross P, Witte U, Gooday AJ, Cowie G, Kitazato H.  2013.  Macrofaunal colonization across the Indian margin oxygen minimum zone. Biogeosciences. 10:7161-7177.   10.5194/bg-10-7161-2013   AbstractWebsite

There is a growing need to understand the ability of bathyal assemblages to recover from disturbance and oxygen stress, as human activities and expanding oxygen minimum zones increasingly affect deep continental margins. The effects of a pronounced oxygen minimum zone (OMZ) on slope benthic community structure have been studied on every major upwelling margin; however, little is known about the dynamics or resilience of these benthic populations. To examine the influence of oxygen and phytodetritus on shortterm settlement patterns, we conducted colonization experiments at 3 depths on the West Indian continental margin. Four colonization trays were deployed at each depth for 4 days at 542 and 802 m (transect 1-16 degrees 58 ' N) and for 9 days at 817 and 1147 m (transect 2-17 degrees 31 ' N). Oxygen concentrations ranged from 0.9 mu M (0.02 mLL(-1)) at 542 m to 22 mu M (0.5 mLL(-1) ) at 1147 m. All trays contained local defaunated sediments; half of the trays at each depth also contained C-13/N-15-labeled phytodetritus mixed into the sediments. Sediment cores were collected between 535 m and 1140 m from 2 cross-margin transects for analysis of ambient (source) macrofaunal (> 300 mu m) densities and composition. Ambient macrofaunal densities ranged from 0 ind m(-2) (at 535-542 m) to 7400 ind m(-2), with maximum values on both transects at 700-800 m. Macrofaunal colonizer densities ranged from 0 ind m(-2) at 542 m, where oxygen was lowest, to average values of 142 ind m(-2) at 800 m, and 3074 ind m(-2) at 1147 m, where oxygen concentration was highest. These were equal to 4.3 and 151% of the ambient community at 800 m and 1147 m, respectively. Community structure of settlers showed no response to the presence of phytodetritus. Increasing depth and oxygen concentration, however, significantly influenced the community composition and abundance of colonizing macrofauna. Polychaetes constituted 92.4% of the total colonizers, followed by crustaceans (4.2%), mollusks (2.5%), and echinoderms (0.8%). The majority of colonizers were found at 1147 m; 88.5% of these were Capitella sp., although they were rare in the ambient community. Colonists at 800 and 1147 m also included ampharetid, spionid, syllid, lumbrinerid, cirratulid, cossurid and sabellid polychaetes. Consumption of C-13/N-15-labeled phytodetritus was observed for macrofaunal foraminifera (too large to be colonizers) at the 542 and 802/817 m sites, and by metazoan macrofauna mainly at the deepest, better oxygenated sites. Calcareous foraminifera (Uvigerina, Hoeglundina sp.), capitellid polychaetes and cumaceans were among the major phytodetritus consumers. These preliminary experiments suggest that bottom-water oxygen concentrations may strongly influence ecosystem services on continental margins, as reflected in rates of colonization by benthos and colonizer processing of carbon following disturbance. They may also provide a window into future patterns of settlement on the continental slope as the world's oxygen minimum zones expand.

Ewel, KC, Cressa C, Kneib RT, Lake PS, Levin LA, Palmer MA, Snelgrove P, Wall DH.  2001.  Managing critical transition zones. Ecosystems. 4:452-460.   10.1007/s10021-001-0106-0   AbstractWebsite

Ecosystems that function as critical transition zones (CTZs) among terrestrial, freshwater, and marine habitats are closely connected to the ecosystems adjacent to them and are characterized by a rapid flux of materials and organisms. CTZs play various roles, including mediating water flows, accumulating sediments and organic matter, processing nutrients, and providing opportunities for recreation. They are particularly difficult to manage because they tend to be small, albeit important, components of large watersheds, and managers may not have control over the entire landscape. Moreover, they are often the focus of intensive human activity. Consequently, CTZs are critically important zones, and their preservation and protection are likely to require unique collaboration among scientists, managers, and stakeholders. Scientists can learn a great deal from the study of these ecosystems, taking advantage of small size and the importance of fluxes, but a good understanding of adaptive management strategies is needed to establish a dialogue with managers and stakeholders on technical and management issues. An understanding of risk analysis is also important to help set meaningful goals and establish logical strategies that include all of the interested parties. Successful restoration of a CTZ is the best test of the quality of knowledge about its structure and function. Much has already been learned about coastal CTZs through restoration projects, and the large number of such projects involving riparian CTZs in particular suggests that there is considerable opportunity for fruitful collaborations between scientists and managers.

Grupe, BM, Krach ML, Pasulka AL, Maloney JM, Levin LA, Frieder CA.  2015.  Methane seep ecosystem functions and services from a recently discovered southern California seep. Marine Ecology-an Evolutionary Perspective. 36:91-108.   10.1111/maec.12243   AbstractWebsite

The recent discovery of a methane seep with diverse microhabitats and abundant groundfish in the San Diego Trough (1020 m) off the coast of Del Mar, California raised questions about the role of seep ecosystem functions and services in relation to continental margins. We used multicorer and ROV grab samples and an ROV survey to characterize macrofaunal structure, diversity, and trophic patterns in soft sediments and authigenic carbonates; seep microhabitats and taxa observed; and the abundance and spatial patterns of fishery-relevant species. Biogenic microhabitats near the Del Mar Seep included microbially precipitated carbonate boulders, bacterial mats, vesicomyid clam beds, frenulate and ampharetid beds, vestimentiferan tubeworm clumps, and fields of Bathysiphon filiformis tubes. Macrofaunal abundance increased and mean faunal delta C-13 signatures decreased in multicorer samples nearer the seep, suggesting that chemosynthetic production enhanced animal densities outside the seep center. Polychaetes dominated sediments, and ampharetids became especially abundant near microbial mats, while gastropods, hydroids, and sponges dominated carbonate rocks. A wide range of stable isotopic signatures reflected the diversity of microhabitats, and methane-derived carbon was the most prevalent source of nutrition for several taxa, especially those associated with carbonates. Megafaunal species living near the seep included longspine thornyhead (Sebastolobus altivelis), Pacific dover sole (Microstomus pacificus), and lithodid crabs (Paralomis verrilli), which represent targets for demersal fisheries. Sebastolobus altivelis was especially abundant (6.5-8.2 fish.100 m(-2)) and appeared to aggregate near the most active seep microhabitats. The Del Mar Methane Seep, like many others along the world's continental margins, exhibits diverse ecosystem functions and enhances regional diversity. Seeps such as this one may also contribute ecosystem services if they provide habitat for fishery species, export production to support margin food webs, and serve as sinks for methane-derived carbon.

Thurber, AR, Levin LA, Rowden AA, Sommer S, Linke P, Kroger K.  2013.  Microbes, macrofauna, and methane: A novel seep community fueled by aerobic methanotrophy. Limnology and Oceanography. 58:1640-1656.   10.4319/lo.2013.58.5.1640   AbstractWebsite

During the discovery and description of seven New Zealand methane seep sites, an infaunal assemblage dominated by ampharetid polychaetes was found in association with high seabed methane emission. This ampharetid-bed assemblage had a mean density of 57,000 +/- 7800 macrofaunal individuals m(-2) and a maximum wet biomass of 274 g m(-2), both being among the greatest recorded from deep-sea methane seeps. We investigated these questions: Does the species assemblage present within these ampharetid beds form a distinct seep community on the New Zealand margin? and What type of chemoautotrophic microbes fuel this heterotrophic community? Unlike the other macro-infaunal assemblages, the ampharetid-bed assemblage composition was homogeneous, independent of location. Based on a mixing model of species-specific mass and isotopic composition, combined with published respiration measurements, we estimated that this community consumes 29-90 mmol C m(-2) d(-1) of methane-fueled biomass; this is > 290 times the carbon fixed by anaerobic methane oxidizers in these ampharetid beds. A fatty acid biomarker approach supported the finding that this community, unlike those previously known, consumes primarily aerobic methanotrophic bacteria. Due to the novel microbial fueling and high methane flux rates, New Zealand's ampharetid beds provide a model system to study the influence of metazoan grazing on microbially mediated biogeochemical cycles, including those that involve greenhouse gas emissions.

Kayen, RE, Schwab WC, Lee HJ, Torresan ME, Hein JR, Quinterno PJ, Levin LA.  1989.  Morphology of sea-floor landslides on Horizon Guyot: application of steady-state geotechnical analysis. Deep-Sea Research Part a-Oceanographic Research Papers. 36:1817-1839.   10.1016/0198-0149(89)90114-3   AbstractWebsite

Mass movement and erosion have been identified on the pelagic sediment cap of Horizon Guyot, a seamount in the Mid-Pacific Mountains. Trends in the size, shape and preservation of bedforms and sediment textural trends on the pelagic cap indicate that bottom-current-generated sediment transport direction is upslope. Slumping of the sediment cap occurred on and that the net bedload transport direction is upslope. Slumping of the sediment cap occurred on the northwest side of the guyot on a 1.6° to 2.0° slope in the zone of enhanced bottom-current activity. Submersible investigations of these slump blocks show them to be discrete and to have a relief of 6–15 m, with nodular chert beds cropping out along the headwall of individual rotated blocks. An evaluation of the stability of the sediment cap suggests that the combination of the current-induced beveling of the sea floor and infrequent earthquake loading accompanied by cyclic strength reduction is responsible for the initiation of slumps. The sediment in the area of slumping moved short distances in relatively coherent masses, whereas sediment that has moved beyond the summit cap perimeter has fully mobilized into sediment gravity flows and traveled large distances. A steady-state geotechnical analysis of Horizon Guyot sediment indicates the predisposition of deeply buried sediment towards disintegrative flow failure on appropriately steep slopes. Thus, slope failure in this deeper zone would include large amounts of internal deformation. However, gravitational stress in the near-surface sediment of the summit cap (sub-bottom depth< 14 m) is insufficient to maintain downslope movement after initial failure occurs. The predicted morphology of coherent slump blocks displaced and rafted upon a weakened zone at depth corresponds well with seismic-reflection data and submersible observations.

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

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

Sperling, EA, Frieder CA, Raman AV, Girguis PR, Levin LA, Knoll AH.  2013.  Oxygen, ecology, and the Cambrian radiation of animals. Proceedings of the National Academy of Sciences of the United States of America. 110:13446-13451.   10.1073/pnas.1312778110   AbstractWebsite

The Proterozoic-Cambrian transition records the appearance of essentially all animal body plans (phyla), yet to date no single hypothesis adequately explains both the timing of the event and the evident increase in diversity and disparity. Ecological triggers focused on escalatory predator-prey "arms races" can explain the evolutionary pattern but not its timing, whereas environmental triggers, particularly ocean/atmosphere oxygenation, do the reverse. Using modern oxygen minimum zones as an analog for Proterozoic oceans, we explore the effect of low oxygen levels on the feeding ecology of polychaetes, the dominant macrofaunal animals in deep-sea sediments. Here we show that low oxygen is clearly linked to low proportions of carnivores in a community and low diversity of carnivorous taxa, whereas higher oxygen levels support more complex food webs. The recognition of a physiological control on carnivory therefore links environmental triggers and ecological drivers, providing an integrated explanation for both the pattern and timing of Cambrian animal radiation.

Wishner, KF, Ashjian CJ, Gelfman C, Gowing MM, Kann L, Levin LA, Mullineaux LS, Saltzman J.  1995.  Pelagic and benthic ecology of the lower interface of the Eastern Tropical Pacific oxygen minimum zone. Deep-Sea Research Part I-Oceanographic Research Papers. 42:93-115.   10.1016/0967-0637(94)00021-j   AbstractWebsite

The distributions of pelagic and benthic fauna were examined in relation to the lower boundary of the oxygen minimum zone (OMZ) on and near Volcano 7, a seamount that penetrates this feature in the Eastern Tropical Pacific. Although the broad, pronounced OMZ in this region is an effective barrier for most zooplankton, zooplankton abundances, zooplankton feeding rates, and ambient suspended particulate organic carbon (POC) peaked sharply in the lower OMZ (about 740-800 m), in association with the minimum oxygen concentration and the increasing oxygen levels just below it. Zooplankton in the lower OMZ were also larger in size, and the pelagic community included some very abundant, possibly opportunistic, species. Elevated POC and scatter in the light transmission data suggested the existence of a thin, particle-rich, and carbon-rich pelagic layer at the base of the OMZ. Gut contents of planktonic detritivores implied opportunistic ingestion of bacterial aggregates, possibly from this layer. Benthic megafaunal abundances on the seamount, which extended up to 730 m, peaked at about 800 m. There was a consistent vertical progression in the depth of first occurrence of different megafaunal taxa in this depth range, similar to intertidal zonation. Although the vertical gradients of temperature, salinity, and oxygen were gradual at the lower OMZ interface (in contrast to the upper OMZ interface at the thermocline), temporal variability in oxygen levels due to internal wave-induced vertical excursions of the OMZ may produce the distinct zonation in the benthic fauna. The characteristics of the lower OMZ interface result from biological interactions with the chemical and organic matter gradients of the OMZ. Most zooplankton are probably excluded physiologically from pronounced OMZs. The zooplankton abundance peak at the lower interface of the OMZ occurs where oxygen becomes sufficiently high to permit the zooplankton to utilize the high concentrations of organic particles that have descended through the OMZ relatively unaltered because of low metazoan abundance. A similar scenario applies to megabenthic distributions. Plankton layers and a potential short food chain (bacteria to zooplankton) at OMZ interfaces suggest intense utilization and modification of organic material, localized within a thin midwater depth zone. This could be a potentially significant filter for organic material sinking to the deep-sea floor.

Mehring, AS, Hatt BE, Kraikittikun D, Orelo BD, Rippy MA, Grant SB, Gonzalez JP, Jiang SC, Ambrose RF, Levin LA.  2016.  Soil invertebrates in Australian rain gardens and their potential roles in storage and processing of nitrogen. Ecological Engineering. 97:138-143.   10.1016/j.ecoleng.2016.09.005   AbstractWebsite

Research on rain gardens generally focuses on hydrology, geochemistry, and vegetation. The role of soil invertebrates has largely been overlooked, despite their well-known impacts on soil nutrient storage, removal, and processing. Surveys of three rain gardens in Melbourne, Australia, revealed a soil invertebrate community structure that differed significantly among sites but was stable across sampling dates (July 2013 and April 2014). Megadrilacea (earthworms), Enchytraeidae (potworms), and Collembola (springtails) were abundant in all sites, and together accounted for a median of 80% of total soil invertebrate abundance. Earthworms were positively correlated to soil organic matter content, but the abundances of other taxonomic groups were not strongly related to organic matter content, plant cover, or root biomass across sites. While less than 5% of total soil N was estimated to be stored in the body tissues of these three taxa, and estimated N gas emissions from earthworms (N2O and N-2) were low, ingestion and processing of soil was high (e.g., up to 417% of the upper 5 cm of soil ingested by earthworms annually in one site), suggesting that the contribution of these organisms to N cycling in rain gardens may be substantial. Thus, invertebrate communities represent an overlooked feature of rain garden design that can play an important role in the structure and function of these systems. (C) 2016 Elsevier B.V. All rights reserved.