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Levin, LA, McCann LD, Thomas CL.  1991.  The ecology of polychaetes on deep seamounts in the eastern Pacific Ocean. Ophelia. :467-476. AbstractWebsite

Polychaetes were collected by the submersible ALVIN on 18 deep (788-3,353 m) seamounts in the eastern Pacific Ocean at 10-degrees, 13-degrees, 20-degrees and 30-degrees N off western Mexico. Polychaetes comprised 57.7% of all macrofauna collected. Average density over all locations was 942 polychaetes/m2. Thirty-eight families were represented among the 1,422 infaunal polychaetes collected. Five families, the Paraonidae, Cirratulidae, Syllidae, Ampharetidae, and Sabellidae, attained average densities > 1 individual/196 cm2 core. We evaluated effects of latitude, local setting, depth, and substrate on polychaete abundance, taxonomic composition, and lifestyles. Unusually high polychaete densities (7,194/m2) and low diversities were observed in a shallow caldera (788 m) at 13-degrees N. Excluding this site, the latitude exhibiting the highest polychaete densities (xBAR = 939/m2) was 10-degrees N. Of the seven settings examined, pit craters (within seamount calderas) supported the highest densities (xBAR = 1031/m2), and hydrothermal oxide fields and seamount bases exhibited the lowest polychaete densities (xBAR = 576-612/m2). Rippled foraminiferal sands on volcano summits supported large numbers of filter feeders, particularly sabellids. Regressions of total polychaete abundance on depth and on percent sand were not significant. Large, epifaunal, sediment-agglutinating protozoans (Phylum Sarcodina: Class Xenophyophorea) provided habitat for 34 polychaete species. Polychaete abundance and family composition were generally similar to those reported for other nearshore, deep-sea environments at comparable depths. With the exception of the shallowest site, species richness was typically high.

Levin, LA, Thomas CL.  1988.  The ecology of xenophyophores (Protista) on eastern Pacific seamounts. Deep-Sea Research Part a-Oceanographic Research Papers. 35:2003-2027.   10.1016/0198-0149(88)90122-7   AbstractWebsite

Large, agglutinating protozoans of the class Xenophyophorea are the dominant epifaunal organisms on soft and hard substrates of many bathyal seamounts in the eastern Pacific Ocean off Mexico. Observations made with the submersible Alvin and remotely towed camera sleds on 17 seamounts at 31°, 20°, 13° and 10°N revealed more than ten distinct xenophyophore test morphologies. Most of these appear to represent previously undescribed species. Reticulate forms are numerically dominant at 20°, 13° and 10°N. Xenophyophore abundances increase with decreasing latitude, being rare at 30°N, present at densities of 0.1–1.0 m−2 at 20° and 13°N and often exceeding 1.0 m−2 at 10°N, occasionally reaching 10–18 m−2. Highest concentrations are observed on caldera floors near the base of steep caldera walls, at depths between 1700 and 2500 m. Most individuals select sand-size pelagic foraminiferan tests (63–500 μm) and exclude pebble, silt and clay-size particles for test construction.Xenophyophore on seamounts modify the structure of metazoan communities and may play a role in maintenance of infaunal diversity. Twenty-seven xenophyophore tests were found to provide habitat for 16 major macrofaunal taxa (152 individuals) and three meiofaunal taxa (333 individuals). The presence of xenophyophores also enhances the abundance of isopods, tanaids, ophiuroids, nematodes and harpacticoid copepods dwelling in sediments surrounding the tests. Mobile megafauna are attracted to sediment beneath and adjacent to xenophyophores. We suggest that xenophyophores, which are abundant on many topographic features in deep water (e.g. guyots, trenches, canyons and continental slopes), are a functionally important component of deep-sea benthic communities and require further autecological and synecological investigation.

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.

Levin, LA, Creed EL.  1986.  Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development. Marine Biology. 92:103-113.   10.1007/bf00392752   AbstractWebsite

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.

Levin, LA.  1986.  Effects of enrichment on reproduction in the opportunistic polychaete Streblospio benedicti (Webster): a mesocosm study. Biological Bulletin. 171:143-160.   10.2307/1541913   AbstractWebsite

The influence of organic enrichment on growth and planktotrophic development of the spionid polychaete Streblospio benedicti Webster was examined in two mesocosm experiments conducted at the MERL facility, University of Rhode Island. Specimens of S. benedicti were collected and their reproductive traits monitored near the conclusion of a two-year eutrophication experiment, and in the middle of a sludge addition experiment. Nutrient (N, P, and Si) enrichments at 8× and 32× the average aerial input into Narragansett Bay, Rhode Island, resulted in increases in body length, segment number, and length per segment, and a doubling of brood size in S. benedicti females. These increases were substantially higher during May (12°C) than August (20°C). Enrichment effects were stronger in the 8× than 32× nutrient treatment. In the sewage sludge experiment body size increased 20% over control values at the highest (8×) sludge treatment level (nitrogen loading equivalent to the 8× nutrient treatment) but no significant increase was noted at the 4× sludge level, which received half as much nitrogen as the 8× sludge treatment. Mean brood size increased by a factor of 4.6 over controls in the 8× sludge treatment and by a factor of 2.3 in the 4× sludge treatment. Within the range of adult body sizes observed, brood size enhancement occurred independent of increased length or segment number in both nutrient and sludge enrichment treatments. The ability to translate elevated food supply directly into increased reproductive output may underly opportunistic dynamics in macrobenthos. Brood size enhancement of the magnitude observed probably contributes to the high S. benedicti densities found in polluted or organically enriched settings.

Levin, LA, Demaster DJ, McCann LD, Thomas CL.  1986.  Effects of giant protozoans (Class Xenophyophorea) on deep-seamount benthos. Marine Ecology-Progress Series. 29:99-104.   10.3354/meps029099   AbstractWebsite

Biogenic sediment structures have been proposed to enhance diversity in deep-sea sediments. To evaluate this hypothesis we examined the influence of xenophyophores, giant sediment-agglutinating protozoans, on the structure of metazoan communities inhabiting sediments of deep (1000 to 3300 m) seamounts in the eastern Pacific Ocean. Xenophyophores provided habitat for 16 major metazoan taxa. Sediments immediately surrounding xenophyophores exhibited elevated faunal densities and species richness relative to control sediments collected 1 m from the tests. Amphipods were exclusively associated with the protozoan tests or sediments beneath them. Crustaceans, molluscs, and echinoderms exhibited enhanced infaunal densities in the presence of xenophyophores but polychaetes did not. Both horizontal and vertical distributions of infauna appear to be influenced by these protozoans. 234Th measurements suggest that xenophyophores and their associated fauna increase the particle flux of fine-grained material to the seabed and enhance subsurface mixing on a 100 d time scale. We propose that xenophyophores alter hydrodynamic conditions and provide deep-sea metazoans with substrate, food, and refuge. The resulting habitat heterogeneity may contribute to maintenance of high benthic diversity.

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

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

Bridges, TS, Levin LA, Cabrera D, Plaia G.  1994.  Effects of sediment amended with sewage, algae, or hydrocarbons on growth and reproduction in two opportunistic polychaetes. Journal of Experimental Marine Biology and Ecology. 177:99-119.   10.1016/0022-0981(94)90146-5   AbstractWebsite

The effects of organic- (sewage and blue-green algae) and hydrocarbon- (no. 2 fuel oil) amended sediments on growth schedules, age and size at maturity, fecundity, and reproductive output were examined for the opportunistic polychaetes Streblospio benedicti Webster and Capitella sp. 1. The two species responded very differently to the amended sediments. For S. benedicti, asymptotic size was reduced and age at first reproduction occurred later in the algae and hydrocarbon treatments compared to the marsh mud only and sewage treatments. Organic- and hydrocarbon-amended sediments did not affect per brood measures of fecundity or C and N investment in S. benedicti. In contrast, Capitella sp. I exhibited strong, positive responses to the organically amended sediments; this was seen in terms of more rapid growth (2 x), younger age at first reproduction (50%), larger asymptotic size (6 x), and higher per brood fecundity and C and N investment (4 x). Reproductive output, a relative measure of reproductive investment, was not directly affected by treatments in either species. For Capitella sp. I in organically enriched settings, the benefits of larger body size appear to include higher per brood fecundity without increasing the relative cost (in terms of reproductive output) of producing a brood of young. The population explosions of Capitella sp. I in response to organic enrichment are the result of earlier reproduction and increased body size and fecundity. Positive population-level responses of S. benedicti to contaminated sediments may be the result of its ability to tolerate conditions that other members of a community do not.

Rasmussen, LL, Cornuelle BD, Levin LA, Largier JL, Di Lorenzo E.  2009.  Effects of small-scale features and local wind forcing on tracer dispersion and estimates of population connectivity in a regional scale circulation model. Journal of Geophysical Research-Oceans. 114   10.1029/2008jc004777   AbstractWebsite

A small-scale model of the Southern California-Northern Baja California coastline has been developed to explore dispersion over the continental shelf, with specific attention to physical parameters pertinent to simulations of larval dispersal and population connectivity. The ROMS simulation employs a nested grid system, with an inner domain resolution of 600 m and an outer domain resolution of 1.5 km. Realistic bathymetry and forcing were employed to investigate the effects of passive transport of tracers introduced at locations with known communities of mytilid mussels along the coastline. The effects of topographic resolution, boundary conditions, and choice of meteorological forcing products on dispersion rates, tracer trajectories, and the subsequent measures of population connectivity were examined. In particular, the choice of wind forcing product resulted in different circulation patterns and tracer trajectories and had especially important consequences on measures of larval connectivity such as self-seeding, potential for larval settlement ( import), and contribution to the pool of available larvae ( export). While some forcing products performed better when model data were compared to field measurements, no product was clearly superior. The uncertainty in results, which may appear minor in larger-scale temperature or surface velocity fields, is significant when examining a sensitive passive tracer. This modeling uncertainty needs to be addressed when interpreting connectivity results.

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, [], [URL:] 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.

Levin, LA, Etter RJ, Rex MA, Gooday AJ, Smith CR, Pineda J, Stuart CT, Hessler RR, Pawson D.  2001.  Environmental influences on regional deep-sea species diversity. Annual Review of Ecology and Systematics. 32:51-93.   10.1146/annurev.ecolsys.32.081501.114002   AbstractWebsite

Most of our knowledge of biodiversity and its causes in the deep-sea benthos derives from regional-scale sampling studies of the macrofauna. Improved sampling methods and the expansion of investigations into a wide variety of habitats have revolutionized our understanding of the deep sea. Local species diversity shows clear geographic variation on spatial scales of 100-1000 km. Recent sampling programs have revealed unexpected complexity in community structure at the landscape level that is associated with large-scale oceanographic processes and their environmental consequences. We review the relationships between variation in local species diversity and the regional-scale phenomena of boundary constraints, gradients of productivity, sediment heterogeneity, oxygen availability, hydrodynamic regimes, and catastrophic physical disturbance. We present a conceptual model of how these interdependent environmental factors shape regional-scale variation in local diversity. Local communities in the deep sea may be composed of species that exist as metapopulations whose regional distribution depends on a balance among global-scale, landscape-scale, and small-scale dynamics. Environmental gradients may form geographic patterns of diversity by influencing local processes such as predation, resource partitioning, competitive exclusion, and facilitation that determine species coexistence. The measurement of deep-sea species diversity remains a vital issue in comparing geographic patterns and evaluating their potential causes. Recent assessments of diversity using species accumulation curves with randomly pooled samples confirm the often-disputed claim that the deep sea supports higher diversity than the continental shelf. However, more intensive quantitative sampling is required to fully characterize the diversity of deep-sea sediments, the most extensive habitat on Earth. Once considered to be constant, spatially uniform, and isolated, deep-sea sediments are now recognized as a dynamic, richly textured environment that is inextricably linked to the global biosphere. Regional studies of the last two decades provide the empirical background necessary to formulate and test specific hypotheses of causality by controlled sampling designs and experimental approaches.

Navarro, MO, Bockmon EE, Frieder CA, Gonzalez JP, Levin LA.  2014.  Environmental pH, O-2 and capsular effects on the geochemical composition of statoliths of embryonic squid Doryteuthis opalescens. Water. 6:2233-2254.   10.3390/w6082233   AbstractWebsite

Spawning market squid lay embryo capsules on the seafloor of the continental shelf of the California Current System (CCS), where ocean acidification, deoxygenation and intensified upwelling lower the pH and [O-2]. Squid statolith geochemistry has been shown to reflect the squid's environment (e. g., seawater temperature and elemental concentration). We used real-world environmental levels of pH and [O-2] observed on squid-embryo beds to test in the laboratory whether or not squid statolith geochemistry reflects environmental pH and [O-2]. We asked whether pH and [O-2] levels might affect the incorporation of element ratios (B:Ca, Mg:Ca, Sr:Ca, Ba:Ca, Pb:Ca, U:Ca) into squid embryonic statoliths as (1) individual elements and/or (2) multivariate elemental signatures, and consider future applications as proxies for pH and [O-2] exposure. Embryo exposure to high and low pH and [O-2] alone and together during development over four weeks only moderately affected elemental concentrations of the statoliths, and uranium was an important element driving these differences. Uranium: Ca was eight-times higher in statoliths exposed to low pHT (7.57-7.58) and low [O-2] (79-82 mu than those exposed to higher ambient pHT (7.92-7.94) and [O-2] (241-243 mu In a separate experiment, exposure to low pHT (7.55-7.56) or low [O-2] (83-86 mu yielded elevated U:Ca and Sr:Ca in the low [O-2] treatment only. We found capsular effects on multiple elements in statoliths of all treatments. The multivariate elemental signatures of embryonic statoliths were distinct among capsules, but did not reflect environmental factors (pH and/or [O-2]). We show that statoliths of squid embryos developing inside capsules have the potential to reflect environmental pH and [O-2], but that these "signals" are generated in concert with the physiological effects of the capsules and embryos themselves.

Levin, LA, Childers SE, Smith CR.  1991.  Epibenthic, agglutinating foraminiferans in the Santa Catalina Basin and their response to disturbance. Deep-Sea Research Part a-Oceanographic Research Papers. 38:465-483.   10.1016/0198-0149(91)90047-j   AbstractWebsite

There are five common species of large (0.5-6 cm long) epibenthic, agglutinating foraminiferans in the Santa Catalina Basin (1200-1350 m). This paper describes their basic ecology and response to mound disturbance. Combined, the five species attain mean densities of 200-300 individuals per m2 and their protoplasm has an average biomass of 199.5 mg m-2. Individual species occur at densities ranging from 7 to 100 m-2, and each species has a different population size structure. Protoplasm comprises < 2% of test volumes. Analysis of excess Th-234 revealed no indication of particle sequestering within tests, and acridine orange direct counts of bacteria provided no evidence of microbial gardening or enhancement associated with tests. Twenty-five per cent of tests examined had metazoan associates; approximately half of these were polychaetes. Experiments were carried out to investigate the response of the epibenthic foraminiferal assemblage to disturbance from large, biogenic mounds, a common feature on the Santa Catalina Basin floor. Three branched forms, Pelosina cf. arborescens, P. cf. cylindrica and a mud-walled astrorhizinid, were most abundant on background sediments, less common on natural mounds and absent from artificially-created mounds exposed for 10.5 months. Two spherical species, Oryctoderma sp. and a different mud-walled astrorhizinid, were present at similar densities on artificial mounds (9.5-10.5 months old), natural mounds and undisturbed sediments, but Oryctoderma sp. attained largest sizes on mounds. These two species appear to be opportunistic taxa that can colonize and grow rapidly on mound sediments. This study suggests that disturbance, in this case that by sediment mound builders, is an important source of spatial heterogeneity in deep-water foraminiferal communities. Where sediment mounds occur, foraminiferal assemblages will experience disequilibrium dynamics.

Navarro, MO, Parnell PE, Levin LA.  2018.  Essential market squid (Doryteuthis opalescens) embryo habitat: A baseline for anticipated ocean climate change. Journal of Shellfish Research. 37:601-614.   10.2983/035.037.0313   AbstractWebsite

The market squid Doryteuthis opalescens deposits embryo capsules onto the continental shelf from Baja California to southern Alaska, yet little is known about the environment of embryo habitat. This study provides a baseline of environmental data and insights on factors underlying site selection for embryo deposition off southern California, and defines current essential embryo habitat using (1) remotely operated vehicle-supported surveys of benthos and environmental variables, (2) SCUBA surveys, and (3) bottom measurements of T, S, pH, and O-2. Here, embryo habitat is defined using embryo capsule density, capsule bed area, consistent bed footprint, and association with [O-2] and pH (pCO(2)) on the shelf. Spatial variation in embryo capsule density and location appears dependent on environmental conditions, whereas the temporal pattern of year-round spawning is not. Embryos require [O-2] greater than 160 mu mol and pH(T) greater than 7.8. Temperature does not appear to be limiting (range: 9.9 degrees C-15.5 degrees C). Dense embryo beds were observed infrequently, whereas low-density cryptic aggregations were common. Observations of dense embryo aggregation in response to shoaling of low [O-2] and pH indicate habitat compression. Essential embryo habitat likely expands and contracts in space and time directly with regional occurrence of appropriate O-2 and pH exposure. Embryo habitat will likely be at future risk of compression given secular trends of deoxygenation and acidification within the Southern California Bight. Increasingly localized and dense spawning may become more common, resulting in potentially important changes in market squid ecology and management.

Carson, HS, Cook GS, Lopez-Duarte PC, Levin LA.  2011.  Evaluating the importance of demographic connectivity in a marine metapopulation. Ecology. 92:1972-1984. AbstractWebsite

Recently researchers have gone to great lengths to measure marine metapopulation connectivity via tagging, genetic, and trace-elemental fingerprinting studies. These empirical estimates of larval dispersal are key to assessing the significance of metapopulation connectivity within a demographic context, but the life-history data required to do this are rarely available. To evaluate the demographic consequences of connectivity we constructed seasonal, size-structured metapopulation matrix models for two species of mytilid mussel in San Diego County, California, USA. The self-recruitment and larval exchange terms were produced from a time series of realized connectivities derived from trace-elemental fingerprinting of larval shells during spring and fall from 2003 to 2008. Both species exhibited a strong seasonal pattern of southward movement of recruits in spring and northward movement in fall. Growth and mortality terms were estimated using mark recapture data from representative sites for each species and subpopulation, and literature estimates of juvenile mortality. Fecundity terms were estimated using county-wide settlement data from 2006-2008; these data reveal peak reproduction and recruitment in fall for Mytilus californianus, and spring for M. galloprovincialis. Elasticity and life-stage simulation analyses were employed to identify the season- and subpopulation-specific vital rates and connectivity terms to which the metapopulation growth rate (lambda) was most sensitive. For both species, metapopulation growth was most sensitive to proportional changes in adult fecundity, survival and growth of juvenile stages, and population connectivity, in order of importance, but relatively insensitive to adult growth or survival. The metapopulation concept was deemed appropriate for both Mytilus species as exchange between the subpopulations was necessary for subpopulation persistence. However, highest metapopulation growth occurred in years when a greater proportion of recruits was retained within the predominant source subpopulation. Despite differences in habitat and planktonic duration, both species exhibited similar overall metapopulation dynamics with respect to key life stages and processes. However, different peak reproductive periods in an environment of seasonal current reversals led to different regional (subpopulation) contributions to metapopulation maintenance; this result emphasizes the importance of connectivity analysis for spatial, management of coastal resources.

Sato, KN, Powell J, Rudie D, Levin LA.  2018.  Evaluating the promise and pitfalls of a potential climate change-tolerant sea urchin fishery in southern California. Ices Journal of Marine Science. 75:1029-1041.   10.1093/icesjms/fsx225   AbstractWebsite

Marine fishery stakeholders are beginning to consider and implement adaptation strategies in the face of growing consumer demand and potential deleterious climate change impacts such as ocean warming, ocean acidification, and deoxygenation. This study investigates the potential for development of a novel climate change-tolerant sea urchin fishery in southern California based on Strongylocentrotus fragilis (pink sea urchin), a deep-sea species whose peak density was found to coincide with a current trap-based spot prawn fishery (Pandalus platyceros) in the 200-300-m depth range. Here we outline potential criteria for a climate change-tolerant fishery by examining the distribution, life-history attributes, and marketable qualities of S. fragilis in southern California. We provide evidence of seasonality of gonad production and demonstrate that peak gonad production occurs in the winter season. S. fragilis likely spawns in the spring season as evidenced by consistent minimum gonad indices in the spring/summer seasons across 4 years of sampling (2012-2016). The resiliency of S. fragilis to predicted future increases in acidity and decreases in oxygen was supported by high species abundance, albeit reduced relative growth rate estimates at water depths (485-510 m) subject to low oxygen (11.7-16.9 mmol kg similar to 1) and pHTotal (< 7.44), which may provide assurances to stakeholders and managers regarding the suitability of this species for commercial exploitation. Some food quality properties of the S. fragilis roe (e. g. colour, texture) were comparable with those of the commercially exploited shallow-water red sea urchin (Mesocentrotus franciscanus), while other qualities (e. g. 80% reduced gonad size by weight) limit the potential future marketability of S. fragilis. This case study highlights the potential future challenges and drawbacks of climate-tolerant fishery development in an attempt to inform future urchin fishery stakeholders.

Depatra, KD, Levin LA.  1989.  Evidence of the passive deposition of meiofauna into fiddler crab burrows. Journal of Experimental Marine Biology and Ecology. 125:173-192.   10.1016/0022-0981(89)90095-6   AbstractWebsite

Small-scale biogenic depressions are known to accumulate suspended particles from the water column. Fiddler crab Uca spp. burrows are prevalent depressions in salt marsh sediments and provide habitats for numerous meiofauna. This research investigated the distribution of meiofauna in fiddler crab burrows in a North Carolina salt marsh and examined the hypothesis that patterns of meiofauna abundance in these burrows are the result of passive accumulation of meiofauna. Burrow sediments contained higher densities of total meiofauna than ambient marsh sediments. The differential collection of meiofauna in large (1.6 cm diameter) vs. small (1.1 cm diameter) burrow mimics (test tubes) placed in the field over one tidal cycle suggested that (a) meiofauna are passively trapped in burrows and (b) burrow morphology may cause differential deposition into burrows. These findings were supported by observations of water flow patterns over large and small burrow mimics in a flume in which dye was entrained 0.5–3.5 cm into tubes. Entrainment of meiofauna appeared more likely at flow speeds of 4.3 than 2.5 or 8.7cm·s−1. Adult copepods, foraminifera, and adult and larval polychaetes were equally abundant in natural Uca burrows at all depths sampled. Nematodes, ostracods, juvenile copepods, copepod and noncopepod nauplii, and turbellarians were less abundant in deep (3–16 cm) than shallow (<3 cm) burrow sediments.

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.

Blankenship, LE, Levin LA.  2007.  Extreme food webs: Foraging strategies and diets of scavenging amphipods from the ocean's deepest 5 kilometers. Limnology and Oceanography. 52:1685-1697.   10.4319/lo.2007.52.4.1685   AbstractWebsite

We explore hypotheses that alternate foraging strategies, diet, or nutrient partitioning could help explain the success of scavenging Lysianassoids (Amphipoda) in hadal oligotrophic trenches (depths of 6-11 km) by examining the nutritional strategies of four lysianassoid species ( Eurythenes gryllus, Scopelocheirus schellenbergi, Hirondellea dubia, and Uristes sp. nov.) collected with baited traps (6.3-10.8 km) from the oligotrophic Tonga and Kermadec Trenches (southwest Pacific Ocean). Diets and foraging strategies were examined by use of (1) the nascent DNA-based analysis of hindgut contents, which provides a 'snapshot' of recently ingested organisms, and (2) natural abundance isotopic signatures, which reflect the source of nutrition and relative trophic position. The scavenging guild exhibits remarkable trophic plasticity, and each amphipod species employs alternate foraging modes, including detrivory or predation, to supplement necrophagy. The nutritional strategies of some species appear to shift with age, depth, and even between trenches. Thus, there is no single ubiquitous hadal food web; rather it is influenced by depth and overlying surface productivity. Isotopic data suggest that coexisting species partition the dietary items, providing evidence of competition among members of the scavenging guild. The extreme foraging flexibility of scavenging amphipods may ultimately contribute to their success in severely food-limited hadal ecosystems.

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.

Gooday, AJ, Levin LA, da Silva AA, Bett BJ, Cowie GL, Dissard D, Gage JD, Hughes DJ, Jeffreys R, Lamont PA, Larkin KE, Murty SJ, Schumacher S, Whitcraft C, Woulds C.  2009.  Faunal responses to oxygen gradients on the Pakistan margin: A comparison of foraminiferans, macrofauna and megafauna. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 56:488-502.   10.1016/j.dsr2.2008.10.003   AbstractWebsite

The Pakistan Margin is characterised by a strong mid-water oxygen minimum zone (OMZ) that intercepts the seabed at bathyal depths (150-1300 m). We investigated whether faunal abundance and diversity trends were similar among protists (foraminiferans and gromiids), metazoan macrofauna and megafauna along a transect (140-1850 m water depth) across the OMZ during the 2003 intermonsoon (March-May) and late/post-monsoon (August-October) seasons. All groups exhibited some drop in abundance in the OMZ core (250-500 m water depth; O(2): 0.10-0.13 mL L(-1) = 4.46-5.80 mu M) but to differing degrees. Densities of foraminiferans >63 mu m were slightly depressed at 300 m, peaked at 738 m, and were much lower at deeper stations. Foraminiferans >300 mu m were the overwhelmingly dominant macrofaunal organisms in the OMZ core. Macrofaunal metazoans reached maximum densities at 140 m depth, with additional peaks at 850, 940 and 1850 m where foraminiferans were less abundant. The polychaete Linopherus sp. was responsible for a macrofaunal biomass peak at 950 m. Apart from large swimming animals (fish and natant decapods), metazoan megafauna were absent between 300 and 900 m (O(2) <0.14-0.15 mLL(-1) = 6.25-6.69 mu M) but were represented by a huge, ophiuroid-dominated abundance peak at 1000 m (O(2) similar to 0.15-0.18 mLL(-1) = 6.69-8.03 mu M). Gromiid protists were confined largely to depths below 1150 m (O(2) > 0.2 mLL(-1) = 8.92 mu M). The progressively deeper abundance peaks for foraminiferans (> 63 mu m), Linopherus sp. and ophiuroids probably represent lower OMZ boundary edge effects and suggest a link between body size and tolerance of hypoxia. Macro- and megafaunal organisms collected between 800 and 1100 m were dominated by a succession of different taxa, indicating that the lower part of the OMZ is also a region of rapid faunal change. Species diversity was depressed in all groups in the OMZ core, but this was much more pronounced for macrofauna and megafauna than for foraminiferans. Oxygen levels strongly influenced the taxonomic composition of all faunal groups. Calcareous foraminiferans dominated the seasonally and permanently hypoxic sites (136-300 m); agglutinated foraminiferans were relatively more abundant at deeper stations where oxygen concentrations were >0.13 mLL(-1)( = 5.80 mu M). Polychaetes were the main macrofaunal taxon within the OMZ; calcareous macrofauna, and megafauna (molluscs and echinoderms) were rare or absent where oxygen levels were lowest. The rarity of larger animals between 300 and 700 m on the Pakistan Margin, compared with the abundant macrofauna in the OMZ core off Oman, is the most notable contrast between the two sides of the Arabian Sea. This difference probably reflects the slightly higher oxygen levels and better food quality on the western side. (C) 2008 Published by Elsevier Ltd.

AK, S, LA L, HT R, C S.  2013.  Faunal trophic structure at hydrothermal vents on the southern Mohn’s Ridge, Arctic Ocean. Marine Ecology Progress Series. 473:115-131.   10.3354/meps10050   AbstractWebsite

We explore the trophic ecology of heterotrophic fauna associated with a high temperature (HT) vent, 2 low temperature vents, a ‘near-HT vent’ habitat and a non-vent site situated at upper bathyal depths on the southern Mohn’s Ridge in the Arctic Ocean. Only a single taxon (the gastropod Pseudosetia griegi) was found at the high temperature vent habitat. Their mean δ13C values were significantly lighter than conspecifics from a low temperature vent habitat within the same vent field, reflecting the incorporation of sulfide oxidizing bacteria into the biomass of the animals. The majority of fauna from the low temperature, near-HT, and non-vent habitats had isotopic signatures indicative of assimilation of photosynthetic material. However, we found remarkably diverse isotopic compositions among the fauna sampled here, with a small sub-set of fauna at each site possessing C and N isotopic signatures indicative of incorporation of chemosynthetic production. Moreover, when isotopic signatures of similar taxa were compared from the same sample, δ13C signatures suggested a high degree of trophic complexity can exist over relatively small spatial scales at vent habitats on the southern Mohn’s Ridge. The high contribution of photosynthetic food material to faunal diets and variability in food types may result from the upper bathyal venting depth and sedimentary nature of the vents. We hypothesize that the upper bathyal depth of active venting may lead to iron enhancement of surface photosynthetic production, especially in high nutrient areas, which ultimately sinks to the seabed where it is incorporated by fauna around the vents.

Gallo, ND, Levin LA.  2016.  Fish ecology and evolution in the world's oxygen minimum zones and implications of ocean deoxygenation. Advances in Marine Biology, Vol 74. 74( Curry BE, Ed.).:117-198., San Diego: Elsevier Academic Press Inc   10.1016/bs.amb.2016.04.001   Abstract

Oxygen minimum zones (OMZs) and oxygen limited zones (OLZs) are important oceanographic features in the Pacific, Atlantic, and Indian Ocean, and are characterized by hypoxic conditions that are physiologically challenging for demersal fish. Thickness, depth of the upper boundary, minimum oxygen levels, local temperatures, and diurnal, seasonal, and interannual oxycline variability differ regionally, with the thickest and shallowest OMZs occurring in the subtropics and tropics. Although most fish are not hypoxia-tolerant, at least 77 demersal fish species from 16 orders have evolved physiological, behavioural, and morphological adaptations that allow them to live under the severely hypoxic, hypercapnic, and at times sulphidic conditions found in OMZs. Tolerance to OMZ conditions has evolved multiple times in multiple groups with no single fish family or genus exploiting all OMZs globally. Severely hypoxic conditions in OMZs lead to decreased demersal fish diversity, but fish density trends are variable and dependent on region-specific thresholds. Some OMZ-adapted fish species are more hypoxiatolerant than most megafaunal invertebrates and are present even when most invertebrates are excluded. Expansions and contractions of OMZs in the past have affected fish evolution and diversity. Current patterns of ocean warming are leading to ocean deoxygenation, causing the expansion and shoaling of OMZs, which is expected to decrease demersal fish diversity and alter trophic pathways on affected margins. Habitat compression is expected for hypoxia-intolerant species, causing increased susceptibility to overfishing for fisheries species. Demersal fisheries are likely to be negatively impacted overall by the expansion of OMZs in a warming world.

Nordstrom, MC, Demopoulos AWJ, Whitcraft CR, Rismondo A, McMillan P, Gonzalez JP, Levin LA.  2015.  Food web heterogeneity and succession in created saltmarshes. Journal of Applied Ecology. 52:1343-1354.   10.1111/1365-2664.12473   AbstractWebsite

Ecological restoration must achieve functional as well as structural recovery. Functional metrics for re-establishment of trophic interactions can be used to complement traditional monitoring of structural attributes. In addition, topographic effects on food web structure provide added information within a restoration context; often, created sites may require spatial heterogeneity to effectively match structure and function of natural habitats. We addressed both of these issues in our study of successional development of benthic food web structure, with focus on bottom-up-driven changes in macroinvertebrate consumer assemblages in the saltmarshes of the Venice Lagoon, Italy. We combined quantified estimates of the changing community composition with stable isotope data (C-13:C-12 and N-15:N-14) to compare the general trophic structure between created (2-14years) marshes and reference sites and along topographic elevation gradients within saltmarshes. Macrofaunal invertebrate consumers exhibited local, habitat-specific trophic patterns. Stable isotope-based trophic structure changed with increasing marsh age, in particular with regard to mid-elevation (Salicornia) habitats. In young marshes, the mid-elevation consumer signatures resembled those of unvegetated ponds. The mid-elevation of older and natural marshes had a more distinct Salicornia zone food web, occasionally resembling that of the highest (Sarcocornia-dominated) elevation. In summary, this indicates that primary producers and availability of vascular plant detritus structure consumer trophic interactions and the flow of carbon. Functionally different consumers, subsurface-feeding detritivores (Oligochaeta) and surface grazers (Hydrobia sp.), showed distinct but converging trajectories of isotopic change over time, indicating that successional development may be asymmetric between brown' (detrital) guilds and green' (grazing) guilds in the food web.Synthesis and applications. Created marsh food webs converged into a natural state over about a decade, with successional shifts seen in both consumer community composition and stable isotope space. Strong spatial effects were noted, highlighting the utility of stable isotopes to evaluate functional equivalence in spatially heterogeneous systems. Understanding the recovery of functional properties such as food web support, and their inherent spatial variability, is key to planning and managing successful habitat restoration. Created marsh food webs converged into a natural state over about a decade, with successional shifts seen in both consumer community composition and stable isotope space. Strong spatial effects were noted, highlighting the utility of stable isotopes to evaluate functional equivalence in spatially heterogeneous systems. Understanding the recovery of functional properties such as food web support, and their inherent spatial variability, is key to planning and managing successful habitat restoration.

Gooday, AJ, Bernhard JM, Levin LA, Suhr SB.  2000.  Foraminifera in the Arabian Sea oxygen minimum zone and other oxygen-deficient settings: taxonomic composition, diversity, and relation to metazoan faunas. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 47:25-54.   10.1016/s0967-0645(99)00099-5   AbstractWebsite

Previous work has shown that some foraminiferal species thrive in organically enriched, oxygen-depleted environments. Here, we compare 'live' (stained) faunas in multicorer samples (0-1 cm layer) obtained at two sites on the Oman margin, one located at 412m within the oxygen minimum zone (OMZ) (O(2) = 0.13 ml l(-1)), the other located at 3350 m, well below the main OMZ (O(2) similar to 3.00 ml l(-1)). While earlier studies have focused on the hard-shelled (predominantly calcareous) foraminifera, we consider complete stained assemblages, including poorly known, soft-shelled, monothalamous forms. Densities at the 412-m site were much higher (16,107 individuals.10 cm(-2) in the > 63-mu m fraction) than at the 3350-m site (625 indiv.10 cm(-2)). Species richness (E(S(100))), diversity (H', Fishers Alpha index) and evenness (J') were much lower, and dominance (R1D) was higher, at 412 m compared with 3350 m. At 412 m, small calcareous foraminifera predominated and soft-shelled allogromiids and sacamminids were a minor faunal element. At 3350 m, calcareous individuals were much less common and allogromiids and saccamminids formed a substantial component of the fauna. There were also strong contrasts between the foraminiferal macrofauna( > 300-mu m fraction) at these two sites; relatively small species of Bathysiphon, Globobulimina and Lagenammina dominated at 412 m, very large, tubular, agglutinated species of Bathysiphon, Hyperammina, Rhabdammina and Saccorhiza were important at 3350 m. Our observations suggest that, because they contain fewer soft-shelled and agglutinated foraminifera, a smaller proportion of bathyal, low-oxygen faunas is lost during fossilization compared to faunas from well-oxygenated environments. Trends among foraminifera (> 63 mu m fraction) in the Santa Barbara Basin (590 and 610m depth; O(2) = 0.05 and 0.15 ml(-1) respectively), and macrofaunal foraminifera(> 300 mu m) on the Peru margin (300-1250 m depth: O(2) = 0.02-1.60 mi l(-1)), matched those observed on the Oman margin. Tn particular, soft-shelled monothalamous taxa were rare and large agglutinated taxa were absent in the most oxygen-depleted ( < 0.20 mi l(-1)) stations. Foraminifera often outnumber metazoans (both meiofaunal and macrofaunal) in bathyal oxygen-depleted settings. However, although phylogenetically distant, foraminifera and metazoans exhibit similar population responses to oxygen depletion; species diversity decreases, dominance increases, and the relative abundance of the major taxa changes. The foraminiferal macrofauna ( > 300 mu m) were 5 times more abundant than the metazoan macrofauna at 412 m on the Oman margin but 16 times more abundant at the 3350 m site. Among the meiofauna (63-300 mu m), the trend was reversed, foraminifera were 17 times more abundant than metazoan taxa at 412 m but only 1.4 times more abundant at 3350 In. An abundance of food combined with oxygen levels which are not depressed sufficiently to eliminate the more tolerant taxa, probably explains why foraminifera and macrofaunal metazoans flourished at the 412-m site, perhaps to the detriment of the metazoan meiofauna. (C) 1999 Elsevier Science Ltd. All rights reserved.