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Levin, LA.  2003.  Oxygen minimum zone benthos: Adaptation and community response to hypoxia. Oceanography and Marine Biology, Vol 41. 41:1-45. AbstractWebsite

Mid-water oxygen minima (<0.5ml 1(-1) dissolved O-2) intercept the continental margins along much of the eastern Pacific Ocean, off west Africa and in the Arabian Sea and Bay of Bengal, creating extensive stretches of sea floor exposed to permanent, severe oxygen depletion. These seafloor oxygen minimum zones (OMZs) typically occur at bathyal depths between 200m and 1000m, and are major sites of carbon burial along the continental margins. Despite extreme oxygen depletion, protozoan and metazoan assemblages thrive in these environments. Metazoan adaptations include small, thin bodies, enhanced respiratory surface area, blood pigments such as haemoglobin, biogenic structure formation for stability in soupy sediments, an increased number of pyruvate oxidoreductases, and the presence of sulphide-oxidising symbionts. The organic-rich sediments of these regions often support mats of large sulphide-oxidising bacteria (Thioploca, Beggiatoa, Thiomargarita), and high-density, low-diversity metazoan assemblages. Densities of protistan and metazoan meiofauna are typically elevated in OMZs, probably due to high tolerance of hypoxia, an abundant food supply, and release from predation. Macrofauna and megafauna often exhibit dense aggregations at OMZ edges, but depressed densities and low diversity in the OMZ core, where oxygen concentration is lowest. Taxa most tolerant of severe oxygen depletion (<0.2mll(-1)) in seafloor OMZs include calcareous foraminiferans, nematodes, and annelids. Agglutinated protozoans, harpacticoid copepods, and calcified invertebrates are typically less tolerant. High dominance and relatively low species richness are exhibited by foraminiferans, metazoan meiofauna, and macrofauna within OMZs. At dissolved oxygen concentrations below 0.15 ml l(-1), bioturbation is reduced, the mixed layer is shallow, and chemosynthesis-based nutrition (via heterotrophy and symbiosis) becomes important. OMZs represent a major oceanographic boundary for many species. As they expand and contract over geological time, OMZs may influence genetic diversity and play a key role in the evolution of species at bathyal depths. These ecosystems may preview the types of adaptations, species, and processes that will prevail with increasing hypoxia over ecological and evolutionary time. However, many questions remain unanswered concerning controls on faunal standing stocks in OMZs, and the physiological, enzymatic, metabolic, reproductive and molecular adaptations that permit benthic animals to live in OMZs. As global warming and eutrophication reduce oxygenation of the world ocean, there is a pressing need to understand the functional consequences of oxygen depletion in marine ecosystems.

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.

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Levin, LA.  1994.  Paleoecology and ecology of xenophyophores. Palaios. 9:32-41.   10.2307/3515076   AbstractWebsite

Xenophyophores are large (several mm to 25 cm diameter), agglutinating protozoans, found primarily in the deep sea. Tests range from simple fans, discs or mudballs, to elaborately folded or reticulated forms, and often contain specific particle types or sizes. Xenophyophore densities are highest on sloped sediments associated with seamounts, continental slopes, canyons and trenches, and beneath productive surface waters. Most forms live as epibenthos on hard or soft substrates, feeding on surface deposits and suspended particles, but one endobenthic genus has been described. Modern xenophyophores enhance particle flux to the seabed, creating local regions of intense radiotracer and metazoan activity. Features key to xenophyophore identification in the fossil record include distinct test morphologies that sometimes involve incorporation of globigerinacean tests, sandwich-like structure in cross section, concentrations of barite (found within the protoplasm), and the presence of fecal strands containing many 10 to 20 mm fecal pellets having enhanced Pb contents. While xenophyophores have no confirmed fossil record, modern structures resembling the ichnogenus Paleodictyon are made by endobenthic xenophyophores in the genus Occultammina. Parallel distributions, morphologies and behaviors have been proposed for some graphoglyptid trace fossils, and for some xenophyophores in the families Syringamminidae and Reticulamminidae. Both occur in the deep sea, have regular or irregular network morphologies, and have been proposed to garden bacteria, trap meiofauna, or suspension feed. Fossils previously regarded as phylloid and fucoid algae, and several species of Aschemonella, previously regarded as foraminiferids, have lifestyles and morphologies consistent with those of modern epibenthic xenophyophores. Confirmation of xenophyophore presence in stratigraphic sequences could provide paleohabitat information and help elucidate the origins of this protozoan group.

Levin, LA, Bridges TS.  1995.  Pattern and diversity in reproduction and development. Ecology of marine invertebrate larvae. ( McEdward LR, Ed.).:1-48., Boca Raton: CRC Press Abstract
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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.

Chu, JW, Levin LA.  1989.  Photoperiod and temperature regulation of growth and reproduction in Streblospio benedicti (Polychaeta, Spionidae). Invertebrate Reproduction & Development. 15:131-142. AbstractWebsite

Three experiments were performed to examine responses of the polychaete Streblospio benedicti Webster from North Carolina to combinations of both constant and seasonally changing daylength and temperature. S. benedicti collected in early and late fall were reared under constant (fall) and fall-winter transition daylength and temperature conditions. S. benedicti collected in spring were reared under constant (winter) and winter-spring transition daylength and temperature conditions. In a third experiment, S. benedicti was kept at constant temperature (20°C) and two fixed daylength treatments.Survivorship and maturation of S. benedicti were unaffected by daylength, but body size (setiger number and length), brooding activity and brood size were subject to photoperiod influence. Decreasing daylength and temperature each led to slower growth in the fall-winter transition experiment. Increasing daylength and temperature led to more rapid growth in the winter-spring transition experiment. Long fixed daylength had the same effect. The proportion of females brooding declined under decreasing daylength and/or temperature, but was unaffected by most other treatments. Photoperiod had a significant and unexpected effect on fecundity in all experiments. Brood sizes were larger under conditions of decreasing or increasing daylength relative to fixed treatments. Fixed long daylength produced larger broods of larvae than fixed short daylength. We conclude that photoperiod and temperature act together to regulate growth, reproductive activity and fecundity of S. benedicti and contribute to spring and fall recruitment peaks observed in the field. Episodic events that decouple photoperiod from normal temperature regimes may cause significant fluctuations in reproductive timing and output.

Burkett, AM, Rathburn AE, Perez ME, Levin LA, Cha H, Rouse GW.  2015.  Phylogenetic placement of Cibicidoides wuellerstorfi (Schwager, 1866) from methane seeps and non-seep habitats on the Pacific margin. Geobiology. 13:44-52.   10.1111/gbi.12118   AbstractWebsite

Benthic foraminifera are among the most abundant groups found in deep-sea habitats, including methane seep environments. Unlike many groups, no endemic foraminiferal species have been reported from methane seeps, and to our knowledge, genetic data are currently sparse for Pacific deep-sea foraminifera. In an effort to understand the relationships between seep and non-seep populations of the deep-sea foraminifera Cibicidoides wuellerstorfi, a common paleo-indicator species, specimens from methane seeps in the Pacific were analyzed and compared to one another for genetic similarities of small subunit rDNA (SSU rDNA) sequences. Pacific Ocean C. wuellerstorfi were also compared to those collected from other localities around the world (based on 18S gene available on Genbank, e.g., Schweizer et al., 2009). Results from this study revealed that C. wuellerstorfi living in seeps near Costa Rica and Hydrate Ridge are genetically similar to one another at the species level. Individuals collected from the same location that display opposite coiling directions (dextral and sinstral) had no species level genetic differences. Comparisons of specimens with genetic information available from Genbank (SSU rDNA) showed that Pacific individuals, collected for this study, are genetically similar to those previously analyzed from the North Atlantic and Antarctic. These observations provide strong evidence for the true cosmopolitan nature of C. wuellerstorfi and highlight the importance of understanding how these microscopic organisms are able to maintain sufficient genetic exchange to remain within the same species between seep and non-seep habitats and over global distances.

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

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

McCormick, LR, Levin LA.  2017.  Physiological and ecological implications of ocean deoxygenation for vision in marine organisms. Philosophical Transactions of the Royal Society a-Mathematical Physical and Engineering Sciences. 375   10.1098/rsta.2016.0322   AbstractWebsite

Climate change has induced ocean deoxygenation and exacerbated eutrophication-driven hypoxia in recent decades, affecting the physiology, behaviour and ecology of marine organisms. The high oxygen demand of visual tissues and the known inhibitory effects of hypoxia on human vision raise the questions if and how ocean deoxygenation alters vision in marine organisms. This is particularly important given the rapid loss of oxygen and strong vertical gradients in oxygen concentration in many areas of the ocean. This review evaluates the potential effects of low oxygen (hypoxia) on visual function in marine animals and their implications for marine biota under current and future ocean deoxygenation based on evidence from terrestrial and a few marine organisms. Evolutionary history shows radiation of eye designs during a period of increasing ocean oxygenation. Physiological effects of hypoxia on photoreceptor function and light sensitivity, in combination with morphological changes that may occur throughout ontogeny, have the potential to alter visual behaviour and, subsequently, the ecology of marine organisms, particularly for fish, cephalopods and arthropods with `fast' vision. Visual responses to hypoxia, including greater light requirements, offer an alternative hypothesis for observed habitat compression and shoaling vertical distributions in visual marine species subject to ocean deoxygenation, which merits further investigation. This article is part of the themed issue 'Ocean ventilation and deoxygenation in a warming world'.

Neira, C, Vales M, Mendoza G, Hoh E, Levin LA.  2018.  Polychlorinated biphenyls (PCBs) in recreational marina sediments of San Diego Bay, southern California. Marine Pollution Bulletin. 126:204-214.   10.1016/j.marpolbul.2017.10.096   AbstractWebsite

Polychlorinated biphenyl (PCB) concentrations were determined in surface sediments from three recreational marinas in San Diego Bay, California. Total PCB concentrations ranged from 23 to 153, 31-294, and 151-1387 ng g(-1) for Shelter Island Yacht Basin (SIYB), Harbor Island West (HW) and Harbor Island East (HE), respectively. PCB concentrations were significantly higher in HE and PCB group composition differed relative to HW and SIYB, which were not significantly different from each other in concentration or group composition. In marina sediments there was a predominance (82-85%) of heavier molecular weight PCBs with homologous groups (6CL-7CL) comprising 59% of the total. In HE 75% of the sites exceeded the effect range median (ERM), and toxicity equivalence (TEQ dioxin-like PCBs) values were higher relative to those of HW and SIYB, suggesting a potential ecotoxicological risk.

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

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

Cook, GS, Parnell PE, Levin LA.  2014.  Population connectivity shifts at high frequency within an open-coast marine protected area network. Plos One. 9   10.1371/journal.pone.0103654   AbstractWebsite

A complete understanding of population connectivity via larval dispersal is of great value to the effective design and management of marine protected areas (MPA). However empirical estimates of larval dispersal distance, self-recruitment, and within season variability of population connectivity patterns and their influence on metapopulation structure remain rare. We used high-resolution otolith microchemistry data from the temperate reef fish Hypsypops rubicundus to explore biweekly, seasonal, and annual connectivity patterns in an open-coast MPA network. The three MPAs, spanning 46 km along the southern California coastline were connected by larval dispersal, but the magnitude and direction of connections reversed between 2008 and 2009. Self-recruitment, i.e. spawning, dispersal, and settlement to the same location, was observed at two locations, one of which is a MPA. Self-recruitment to this MPA ranged from 50-84%; within the entire 60 km study region, self-recruitment accounted for 45% of all individuals settling to study reefs. On biweekly time scales we observed directional variability in alongshore current data and larval dispersal trajectories; if viewed in isolation these data suggest the system behaves as a source-sink metapopulation. However aggregate biweekly data over two years reveal a reef network in which H. rubicundus behaves more like a well-mixed metapopulation. As one of the few empirical studies of population connectivity within a temperate open coast reef network, this work can inform the MPA design process, implementation of ecosystem based management plans, and facilitate conservation decisions.

Levin, LA, Gooday AJ.  1992.  Possible roles for xenophyophores in deep-sea carbon cycling. Deep-sea food chains and the global carbon cycle. ( Rowe GT, Pariente V, Eds.).:93-104., Dordrecht ; Boston: Kluwer Academic Publishers Abstract
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Mehring, AS, Levin LA.  2015.  Potential roles of soil fauna in improving the efficiency of rain gardens used as natural stormwater treatment systems. Journal of Applied Ecology. 52:1445-1454.   10.1111/1365-2664.12525   AbstractWebsite

Natural treatment systems such as rain gardens aim to overcome the negative effects of urbanization on water quality, availability, and freshwater and marine ecosystem integrity by mimicking the natural water cycle in urban planning and design. While soils in these systems are inhabited by a diverse array of invertebrates, the soil macrofauna is ignored in the vast majority of studies on new or existing rain gardens. Here, we review the functional roles of invertebrates commonly found within soils of rain gardens. Soil fauna have the potential to substantially alter plant growth, water infiltration rates, and the retention and removal of pathogens, nutrients, heavy metals and other contaminants. Their lack of inclusion in controlled laboratory or greenhouse studies may contribute to differences in observed function in field and laboratory settings. Promising future research directions include the following: (i) the use of controlled experiments to study invertebrate effects on rain garden function; (ii) determining the factors affecting variability in organismal abundance among and within sites; and (iii) the design of rain gardens to facilitate development of fauna that promote desired functions.Synthesis and applications. Soil fauna may substantially alter the function of rain gardens as natural stormwater treatment systems in urban areas. Therefore, incorporat-ing animal effects into design and testing may better enable managers and researchers tounderstand and optimize rain garden functioning, and forecast the longevity of rain gardens. Soil fauna may substantially alter the function of rain gardens as natural stormwater treatment systems in urban areas. Therefore, incorporat-ing animal effects into design and testing may better enable managers and researchers tounderstand and optimize rain garden functioning, and forecast the longevity of rain gardens.

Parker, EA, Rippy MA, Mehring AS, Winfrey BK, Ambrose RF, Levin LA, Grant SB.  2017.  Predictive power of clean bed filtration theory for fecal indicator bacteria removal in stormwater biofilters. Environmental Science & Technology. 51:5703-5712.   10.1021/acs.est.7b00752   AbstractWebsite

Green infrastructure (also referred to as low impact development, or LID) has the potential to transform urban stormwater runoff from an environmental threat to a valuable water resource. In this paper we focus on the removal of fecal indicator bacteria (FIB, a pollutant responsible for runoff associated inland and coastal beach closures) in stormwater biofilters (a common type of green infrastructure). Drawing on a combination of previously published and new laboratory studies of FIB removal in biofilters, we find that 66% of the variance in FIB removal rates can be explained by clean bed filtration theory (CBFT, 31%), antecedent dry period (14%), study effect (8%), biofilter age (7%), and the presence or absence of shrubs (6%). Our analysis suggests that, with the exception of shrubs, plants affect FIB removal indirectly by changing the infiltration rate, not directly by changing the FIB removal mechanisms or altering filtration rates in ways not already accounted for by CBFT. The analysis presented here represents a significant step forward in our understanding of how physicochemical theories (such as CBFT) can be melded with hydrology, engineering design, and ecology to improve the water quality benefits of green infrastructure.

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Levin, L, Blair N, DeMaster D, Plaia G, Fornes W, Martin C, Thomas C.  1997.  Rapid subduction of organic matter by maldanid polychaetes on the North Carolina slope. Journal of Marine Research. 55:595-611.   10.1357/0022240973224337   AbstractWebsite

In situ tracer experiments conducted on the North Carolina continental slope reveal that tube-building worms (Polychaeta: Maldanidae) can, without ingestion, rapidly subduct freshly deposited, algal carbon (C-13-labeled diatoms) and inorganic materials (slope sediment and glass beads) to depths of 10 cm or more in the sediment column. Transport over 1.5 days appears to be nonselective but spatially patchy, creating localized, deep hotspots. As a result of this transport, relatively fresh organic matter becomes available soon after deposition to deep-dwelling microbes and other infauna, and both aerobic and anaerobic processes may be enhanced. Comparison of tracer subduction with estimates from a diffusive mixing model using Th-234-based coefficients, suggests that maldanid subduction activities, within 1.5 d of particle deposition, could account for 25-100% of the mixing below 5 cm that occurs on 100-day time scales. Comparisons of community data from the North Carolina slope for different places and times indicate a correlation between the abundance of deep-dwelling maldanids and the abundance and the dwelling depth in the sediment column of other infauna. Pulsed inputs of organic matter occur frequently in margin environments and maldanid polychaetes are a common component of continental slope macrobenthos. Thus, the activities we observe are likely to be widespread and significant for chemical cycling (natural and anthropogenic materials) on the slope. We propose that species like maldanids, that rapidly redistribute labile organic matter within the seabed, probably function as keystone resource modifiers. They may exert a disproportionately strong influence (relative to their abundance) on the structure of infaunal communities and on the timing, location and nature of organic matter diagenesis and burial in continental margin sediments.

Levin, LA, Huggett D, Myers P, Bridges T, Weaver J.  1993.  Rare earth tagging methods for the study oflarval dispersal by marine invertebrates. Limnology and Oceanography. 38:346-360. AbstractWebsite

Rare-earth elements (REEs) are proposed as a label for invertebrate larvae (clams, barnacles, and polychaetes) and passive larval mimics (polystyrene spheres) to be used in field studies of larval dispersal. Different elements, their uptake levels, retention times, and effects on larval survivorship were examined to assess suitability of REEs as larval tags. Lu, Sm, and Eu were selected for labeling work based on extremely low levels in natural plankton and seawater and due to the high sensitivity of neutron activation analysis to these elements. Adequate label retention occurs on time scales of hours to days in both larvae and microspheres. Combined use of Eu-labeled microspheres and Lu- and Sm-labeled larvae in release-recapture mode offers a highly sensitive method for studying trajectories and behavior of larvae in natural settings. However, dilution factors may limit experimentation to small and intermediate spatial scales (meters to kilometers) in the field.

Levin, LA.  2006.  Recent progress in understanding larval dispersal: new directions and digressions. Integrative and Comparative Biology. 46:282-297.   10.1093/icb/024   AbstractWebsite

Larvae have been difficult to study because their small size limits our ability to understand their behavior and the conditions they experience. Questions about larval transport focus largely on (a) where they go [dispersal] and (b) where they come from [connectivity]. Mechanisms of transport have been intensively studied in recent decades. As our ability to identify larval sources develops, the consequences of connectivity are garnering more consideration. Attention to transport and connectivity issues has increased dramatically in the past decade, fueled by changing motivations that now include management of fisheries resources, understanding of the spread of invasive species, conservation through the design of marine reserves, and prediction of climate-change effects. Current progress involves both technological advances and the integration of disciplines and approaches. This review focuses on insights gained from physical modeling, chemical tracking, and genetic approaches. I consider how new findings are motivating paradigm shifts concerning (1) life-history consequences; (2) the openness of marine populations, self-recruitment, and population connectivity; (3) the role of behavior; and (4) the significance of variability in space and time. A challenge for the future will be to integrate methods that address dispersal on short (intragenerational) timescales such as elemental fingerprinting and numerical simulations with those that reflect longer timescales such as gene flow estimates and demographic modeling. Recognition and treatment of the continuum between ecological and evolutionary timescales will be necessary to advance the mechanistic understanding of larval and population dynamics.

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

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

Whitcraft, CR, Levin LA.  2007.  Regulation of benthic algal and animal communities by salt marsh plants: Impact of shading. Ecology. 88:904-917.   10.1890/05-2074   AbstractWebsite

Plant cover is a fundamental feature of many coastal marine and terrestrial systems and controls the structure of associated animal communities. Both natural and human-mediated changes in plant cover influence abiotic sediment properties and thus have cascading impacts on the biotic community. Using clipping ( structural) and light ( shading) manipulations in two salt marsh vegetation zones ( one dominated by Spartina foliosa and one by Salicornia virginica), we tested whether these plant species exert influence on abiotic environmental factors and examined the mechanisms by which these changes regulate the biotic community. In an unshaded ( plant and shade removal) treatment, marsh soils exhibited harsher physical properties, a microalgal community composition shift toward increased diatom dominance, and altered macrofaunal community composition with lower species richness, a larger proportion of insect larvae, and a smaller proportion of annelids, crustaceans, and oligochaetes compared to shaded ( plant removal, shade mimic) and control treatment plots. Overall, the shaded treatment plots were similar to the controls. Plant cover removal also resulted in parallel shifts in microalgal and macrofaunal isotopic signatures of the most dynamic species. This suggests that animal responses are seen mainly among microalgae grazers and may be mediated by plant modi. cation of microalgae. Results of these experiments demonstrate how light reduction by the vascular plant canopy can control salt marsh sediment communities in an arid climate. This research facilitates understanding of sequential consequences of changing salt marsh plant cover associated with climate or sea level change, habitat degradation, marsh restoration, or plant invasion.

Levin, LA, Gage JD.  1998.  Relationships between oxygen, organic matter and the diversity of bathyal macrofauna. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 45:129-163.   10.1016/s0967-0645(97)00085-4   AbstractWebsite

The relationships of environmental factors with measures of macrobenthic community diversity were examined for the total fauna, and for polychaetes only, from 40 bathyal stations in the North Atlantic, eastern Pacific and Indian Oceans (154-3400 m). Stepwise multiple regression revealed that depth, latitude, sediment organic-carbon content and bottom-water oxygen concentration are significant factors that together explained 52-87% of the variation in macrobenthic species richness (E[s(100)]), the Shannon-Wiener index (H'), dominance (D), and evenness (J'). Percent sand and percent clay were not significant factors. After removal of depth and latitudinal effects, oxygen and organic-carbon concentrations combined accounted for 47, 67, 52 and 32% of residual variation in macrobenthic E(s(100)), H', D, and J', respectively. Organic carbon exhibited a stronger relationship than oxygen to measures of community evenness, and appeared to have more explanatory power for polychaetes than total macrobenthos. When only stations with oxygen < 1mll(-1) were considered, oxygen concentration became the dominant parameter after depth. Results suggest existence of an oxygen threshold ( < 0.45 mi l(-1)), above which oxygen effects on macrobenthic diversity are minor relative to organic matter influence, but below which oxygen becomes a critical factor. Our regression results lead us to hypothesize that for bathyal faunas, oxygen at low concentrations has more influence on species richness, while organic carbon regulates the distribution of individuals among species (community evenness). Examination of rarefaction curves for Indo-Pacific stations reveals that total macrobenthos, polychaetes, crustaceans and molluscs all exhibit reduced species richness within oxygen minimum zones (OMZs). However, representation under conditions of hypoxia varies among taxa, with polychaetes being most tolerant. Molluscs and crustaceans often (but not always) exhibit few individuals and species in OMZs, and sometimes disappear altogether, contributing to reduced macrobenthic diversity and elevated dominance in these settings. The linear negative relationship observed between bathyal species richness and sediment organic-carbon content (used here as a proxy for food availability) may represent the right side (more productive half) of the hump-shaped, diversity-productivity curve reported in other systems. These analyses suggest then are potentially strong influences of organic matter and oxygen on the diversity and composition of bathyal macrobenthos, especially in the Indo-Pacific Ocean. (C) 1998 Elsevier Science Ltd. All rights reserved.

Blair, NE, Plaia GR, Boehme SE, Demaster DJ, Levin LA.  1994.  The remineralization of organic carbon on the North Carolina continental slope. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 41:755-766.   10.1016/0967-0645(94)90046-9   AbstractWebsite

The sources and fates of metabolizable organic carbon were examined at three sites on the North Carolina slope positioned offshore of Cape Fear, Cape Lookout and Cape Hatteras. The C-13/C-12 compositions (delta(13)C) of the solid phase organic matter, and the dissolved inorganic carbon (Sigma CO2) produced during its oxidation, suggested that the labile fraction was predominantly marine in origin. The Sigma CO2 concentration gradient across the sediment-water interface, and by inference the Sigma CO2 flux and production rate, increased northward from Cape Fear to Cape Hatteras. Methane distributions and Sigma CO2 delta(13)C values suggest that the rate of anaerobic diagenesis increased northward as well. The differences in sedimentary biogeochemistry are most likely driven by an along-slope gradient of reactive organic carbon flux to the seabed. This trend in reactive organic carbon flux correlates well with macrofaunal densities previously observed at the three sites. Proximity to the shelf and the transport of particulate material by surface boundary currents may control the deposition of metabolizable material on the Carolina slope. Evidence for methanogenesis was found only on the Cape Hatteras slope. The methane, which was produced at a depth of approximately 1 m in the seabed, was consumed nearly quantitatively in the biologically mixed layer as it diffused upward. Irrigation of the sediments by infauna may have provided the necessary oxidant for the consumption of the methane.

Carson, HS, Lopez-Duarte PC, Rasmussen L, Wang DX, Levin LA.  2010.  Reproductive timing alters population connectivity in marine metapopulations. Current Biology. 20:1926-1931.   10.1016/j.cub.2010.09.057   AbstractWebsite

Populations of most marine organisms are connected by the dispersal of larval stages, with profound implications for marine conservation [1]. Because of the extreme effort needed to empirically measure larval exchange, multispecies conservation efforts must estimate connectivity by extrapolation using taxonomy, adult distribution, life history, behavior, or phenology. Using a 6-year record of connectivity realized through trace-elemental fingerprinting of larval shells, we document the seasonal and interannual variability of larval exchange for two congeneric mussel species with overlapping but distinct distribution, life history, and reproduction timing. We reveal consistent autumn poleward movement and spring equatorward movement for both species, coincident with near-shore surface currents. However, because the major reproductive seasons differ, the dominant source-sink dynamics of these two congeneric species are nearly opposite. Consideration of present and future reproductive timing as altered by climate change is crucial to marine connectivity and conservation, especially for the numerous coastal areas subject to seasonal current reversals.

Levin, LA, Smith CR.  1984.  Response of background fauna to disturbance and enrichment in the deep sea: a sediment tray experiment. Deep-Sea Research Part a-Oceanographic Research Papers. 31:1277-&.   10.1016/0198-0149(84)90001-3   AbstractWebsite

Free-vehicle sediment trays were deployed on the floor of the Santa Catalina Basin (1300 m) to study the response of deep-sea infauna to defaunation and algal enrichment. We recovered three trays that had been filled with frozen basin sediment: one otherwise untreated, one enriched with 20 g of powdered Macrocystis pyrifera (kelp) and containing oxidized sediment, and one kelp-enriched and anoxic. After 4.7 months on the basin floor, macrofaunal density in the unenriched tray was 10% () of that in the surrounding sea floor. Most colonists belonged to species dominant in the background basin community. Juveniles and mature individuals were present, suggesting short maturation times or post-larval dispersal ability in some species. The kelp-enriched (oxidized) tray contained a single macrofaunal mite (Arachnida:Acarina), suggesting avoidance of kelp-enriched sediment by available colonists. The anoxic, kelp-enriched tray contained two dorvilleid polychaetes apparently adapted to high sulphide environments.As in other deep-sea studies, we obtained slower colonization of sediment trays than in shallow water experiments. The primary tray respondents, however, were dominants of the background community, in contrast to previous deep-sea experiments. The findings are consistent with the hypothesis that disturbance is important to the ecology of the dominant species in the Santa Catalina Basin.

Sato, KN, Andersson AJ, Day JMD, Taylor JRA, Frank MB, Jung JY, McKittrick J, Levin LA.  2018.  Response of sea urchin fitness traits to environmental gradients across the Southern California oxygen minimum zone. Frontiers in Marine Science. 5   10.3389/fmars.2018.00258   AbstractWebsite

Marine calcifiers are considered to be among the most vulnerable taxa to climate-forced environmental changes occurring on continental margins with effects hypothesized to occur on microstructural, biomechanical, and geochemical properties of carbonate structures. Natural gradients in temperature, salinity, oxygen, and pH on an upwelling margin combined with the broad depth distribution (100-1,100 m) of the pink fragile sea urchin, Strongylocentrotus (formerly Allocentrotus) fragilis, along the southern California shelf and slope provide an ideal system to evaluate potential effects of multiple climate variables on carbonate structures in situ. We measured, for the first time, trait variability across four distinct depth zones using natural gradients as analogues for species-specific implications of oxygen minimum zone (OMZ) expansion, deoxygenation and ocean acidification. Although S. fragilis may likely be tolerant of future oxygen and pH decreases predicted during the twenty-first century, we determine from adults collected across multiple depth zones that urchin size and potential reproductive fitness (gonad index) are drastically reduced in the OMZ core (450-900 m) compared to adjacent zones. Increases in porosity and mean pore size coupled with decreases in mechanical nanohardness and stiffness of the calcitic endoskeleton in individuals collected from lower pH(Total) (7.57-7.59) and lower dissolved oxygen (13-42 mu mol kg(-1)) environments suggest that S. fragilis may be potentially vulnerable to crushing predators if these conditions become more widespread in the future. In addition, elemental composition indicates that S. fragilis has a skeleton composed of the low Mg-calcite mineral phase of calcium carbonate (mean Mg/Ca = 0.02 mol mol(-1)), with Mg/Ca values measured in the lower end of values reported for sea urchins known to date. Together these findings suggest that ongoing declines in oxygen and pH will likely affect the ecology and fitness of a dominant echinoid on the California margin.