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

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

2018
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.

Neira, C, Ingels J, Mendoza G, Hernandez-Lopez E, Levin LA.  2018.  Distribution of meiofauna in bathyal sediments influenced by the oxygen minimum zone off Costa Rica. Frontiers in Marine Science. 5   10.3389/fmars.2018.00448   AbstractWebsite

Ocean deoxygenation has become a topic of increasing concern because of its potential impacts on marine ecosystems, including oxygen minimum zone (OMZ) expansion and subsequent benthic effects. We investigated the influence of oxygen concentration and organic matter (OM) availability on metazoan meiofauna within and below an OMZ in bathyal sediments off Costa Rica, testing the hypothesis that oxygen and OM levels are reflected in meiofaunal community structures and distribution. Mean total densities in our sampling cores (400-1800 m water depth) were highest with 3688 ind. 10 cm(-2) at the OMZ core at 400 m water depth, decreasing rapidly downslope. Nematodes were overall dominant, with a maximum of 99.9% in the OMZ core, followed by copepods (13%), nauplii (4.8%), and polychaetes (3%). Relative copepod and nauplii abundance increased consistently with depth and increasing bottom-water O-2. Meiofaunal composition was significantly different among sites, with lower taxonomic diversity at OMZ sites relative to deeper, oxygenated sites. Vertical distribution patterns within sediments showed that in strongly oxygen-depleted sites less meiofauna was concentrated in the surface sediment than at deeper slope sites. Highest meiofaunal abundance and lowest diversity occurred under lowest oxygen and highest pigment levels, whereas highest diversity occurred under highest oxygen-concentrations and low pigments, as well as high quality of sedimentary pigment (chl a/phaeo) and organic carbon (C/N). The lower meiofaunal diversity, and lower structural and trophic complexity, at oxygen-depleted sites raises concerns about changes in the structure and function of benthic marine ecosystems in the face of OMZ expansions.

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.

Niner, HJ, Ardron JA, Escobar EG, Gianni M, Jaeckel A, Jones DOB, Levin LA, Smith CR, Thiele T, Turner PJ, Vandover CL, Watling L, Gjerde KM.  2018.  Deep-sea mining with no net loss of biodiversity-an impossible aim. Frontiers in Marine Science. 5   10.3389/fmars.2018.00053   AbstractWebsite

Deep-sea mining is likely to result in biodiversity loss, and the significance of this to ecosystem function is not known. "Out of kind" biodiversity offsets substituting one ecosystem type (e.g., coral reefs) for another (e.g., abyssal nodule fields) have been proposed to compensate for such loss. Here we consider a goal of no net loss (NNL) of biodiversity and explore the challenges of applying this aim to deep seabed mining, based on the associated mitigation hierarchy (avoid, minimize, remediate). We conclude that the industry cannot at present deliver an outcome of NNL. This results from the vulnerable nature of deep-sea environments to mining impacts, currently limited technological capacity to minimize harm, significant gaps in ecological knowledge, and uncertainties of recovery potential of deep-sea ecosystems. Avoidance and minimization of impacts are therefore the only presently viable means of reducing biodiversity losses from seabed mining. Because of these constraints, when and if deep-sea mining proceeds, it must be approached in a precautionary and step-wise manner to integrate new and developing knowledge. Each step should be subject to explicit environmental management goals, monitoring protocols, and binding standards to avoid serious environmental harm and minimize loss of biodiversity. "Out of kind" measures, an option for compensation currently proposed, cannot replicate biodiversity and ecosystem services lost through mining of the deep seabed and thus cannot be considered true offsets. The ecosystem functions provided by deep-sea biodiversity contribute to a wide range of provisioning services (e.g., the exploitation of fish, energy, pharmaceuticals, and cosmetics), play an essential role in regulatory services (e.g., carbon sequestration) and are important culturally. The level of "acceptable" biodiversity loss in the deep sea requires public, transparent, and well-informed consideration, as well as wide agreement. If accepted, further agreement on how to assess residual losses remaining after the robust implementation of the mitigation hierarchy is also imperative. To ameliorate some of the inter-generational inequity caused by mining-associated biodiversity losses, and only after all NNL measures have been used to the fullest extent, potential compensatory actions would need to be focused on measures to improve the knowledge and protection of the deep sea and to demonstrate benefits that will endure for future generations.

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

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

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.

2017
Neira, C, Cossaboon J, Mendoza G, Hoh E, Levin LA.  2017.  Occurrence and distribution of polycyclic aromatic hydrocarbons in surface sediments of San Diego Bay marinas. Marine Pollution Bulletin. 114:466-479.   10.1016/j.marpolbul.2016.10.009   Abstract

Polycyclic aromatic hydrocarbons (PAHs) have garnered much attention due to their bioaccumulation, carcinogenic properties, and persistence in the environment. Investigation of the spatial distribution, composition, and sources of PAHs in sediments of three recreational marinas in San Diego Bay, California revealed significant differences among marinas, with concentrations in one site exceeding 16,000 ng g− 1. ‘Hotspots’ of PAH concentration suggest an association with stormwater outfalls draining into the basins. High-molecular weight PAHs (4–6 rings) were dominant (> 86%); the average percentage of potentially carcinogenic PAHs was high in all sites (61.4–70%) but ecotoxicological risks varied among marinas. Highly toxic benzo(a)pyrene (BaP) was the main contributor (> 90%) to the total toxic equivalent quantity (TEQ) in marinas. PAHs in San Diego Bay marina sediments appear to be derived largely from pyrogenic sources, potentially from combustion products that reach the basins by aerial deposition and stormwater drainage from nearby streets and parking lots.

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

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

Levin, LA, Baco AR, Bowden DA, Colaco A, Cordes EE, Cunha MR, Demopoulos AWJ, Gobin J, Grupe BM, Le J, Metaxas A, Netburn AN, Rouse GW, Thurber AR, Tunnicliffe V, Van Dover CL, Vanreusel A, Watling L.  2016.  Hydrothermal vents and methane seeps: Rethinking the sphere of influence. Frontiers in Marine Science. 3   10.3389/fmars.2016.00072   AbstractWebsite

Although initially viewed as oases within a barren deep ocean, hydrothermal vent and methane seep communities are now recognized to interact with surrounding ecosystems on the sea floor and in the water column, and to affect global geochemical cycles. The importance of understanding these interactions is growing as the potential rises for disturbance from oil and gas extraction, seabed mining and bottom trawling. Here we synthesize current knowledge of the nature, extent and time and space scales of vent and seep interactions with background systems. We document an expanded footprint beyond the site of local venting or seepage with respect to elemental cycling and energy flux, habitat use, trophic interactions, and connectivity. Heat and energy are released, global biogeochemical and elemental cycles are modified, and particulates are transported widely in plumes. Hard and biotic substrates produced at vents and seeps are used by “benthic background” fauna for attachment substrata, shelter, and access to food via grazing or through position in the current, while particulates and fluid fluxes modify planktonic microbial communities. Chemosynthetic production provides nutrition to a host of benthic and planktonic heterotrophic background species through multiple horizontal and vertical transfer pathways assisted by flow, gamete release, animal movements, and succession, but these pathways remain poorly known. Shared species, genera and families indicate that ecological and evolutionary connectivity exists among vents, seeps, organic falls and background communities in the deep sea; the genetic linkages with inactive vents and seeps and background assemblages however, are practically unstudied. The waning of venting or seepage activity generates major transitions in space and time that create links to surrounding ecosystems, often with identifiable ecotones or successional stages. The nature of all these interactions is dependent on water depth, as well as regional oceanography and biodiversity. Many ecosystem services are associated with the interactions and transitions between chemosynthetic and background ecosystems, for example carbon cycling and sequestration, fisheries production, and a host of non-market and cultural services. The quantification of the sphere of influence of vents and seeps could be beneficial to better management of deep-sea environments in the face of growing industrialization.

2015
Neira, C, Mendoza G, Porrachia M, Stransky C, Levin LA.  2015.  Macrofaunal recolonization of copper-contaminated sediments in San Diego Bay. Marine Pollution Bulletin. 101:794-804.   10.1016/j.marpolbul.2015.09.023   AbstractWebsite

Effects of Cu-loading on macrofaunal recolonization were examined in Shelter Island Yacht Basin (San Diego Bay, California). Sediments with high and low Cu levels were defaunated and Cu-spiked, translocated, and then placed back into the environment These demonstrated that the alteration observed in benthic communities associated with Cu contamination occurs during initial recolonization. After a 3-month exposure to sediments with varying Cu levels, two primary colonizing communities were identified: (1) a "mouth assemblage" resembling adjacent background fauna associated with low-Cu levels that was more diverse and predominantly dominated by surface- and subsurface-deposit feeders, burrowers, and tube builders, and (2) a "head assemblage" resembling adjacent background fauna associated with high-Cu concentrations, with few dominant species and an increasing importance of carnivores and mobile epifauna. Cu loading can cause reduced biodiversity and lower structural complexity that may last several months if high concentrations persist, with a direct effect on community functioning. (C) 2015 Elsevier Ltd. All rights reserved.

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.

Raman, AV, Damodaran R, Levin LA, Ganesh T, Rao YKV, Nanduri S, Madhusoodhanan R.  2015.  Macrobenthos relative to the oxygen minimum zone on the East Indian margin, Bay of Bengal. Marine Ecology-an Evolutionary Perspective. 36:679-700.   10.1111/maec.12176   AbstractWebsite

The Bay of Bengal remains one of the least studied of the world's oxygen minimum zones (OMZs). Here we offer a detailed investigation of the macrobenthos relative to oxygen minimum zone [OMZ - DO (dissolved oxygen), concentration <0.5ml1(-1)] at 110 stations off the North East Indian margin (16(0) and 20(0)N) featuring coastal, shelf and slope settings (10-1004m). Macrobenthos (>0.5mm) composition, abundance and diversity were studied in relation to variations in depth, dissolved oxygen, sediment texture and organic carbon. Using multivariate procedures powered by SIMPROF analysis we identified distinct OMZ core sites (depth 150-280m; DO 0.37ml1(-1)) that exhibited dense populations of surface-feeding polychaetes (mean 2188 ind. m(-2)) represented by spionids and cossurids (96%). Molluscs and crustaceans were poorly represented except for ampeliscid amphipods. The lower OMZ sites (DO>0.55mll(-1)) supported a different assemblage of polychaetes (cirratulids, amphinomids, eunicids, orbinids, paraonids), crustaceans and molluscs, albeit with low population densities (mean 343 ind. m(-2)). Species richness [E(S-100)], diversity (Margalef d; H') and evenness (J') were lower and dominance was higher within the OMZ core region. Multiple regression analysis showed that a combination of sand, clay, organic carbon, and dissolved oxygen explained 62-78% of the observed variance in macrobenthos species richness and diversity: E(S-100) and H'. For polychaetes, clay and oxygen proved important. At low oxygen sites (DO <1mll(-1)), depth accounted for most variance. Residual analysis (after removing depth effects) revealed that dissolved oxygen and sediment organic matter influenced 50-62% of residual variation in E(S-100), H' and d for total macrofauna. Of this, oxygen alone influenced up to similar to 50-62%. When only polychaetes were evaluated, oxygen and organic matter explained up to 58-63%. For low oxygen sites, organic matter alone had the explanatory power when dominance among polychaetes was considered. Overall, macrobenthic patterns in the Bay of Bengal were consistent with those reported for other upwelling margins. However, the compression of faunal gradients at shallower depths was most similar to the Chile/Peru margin, and different from the Arabian Sea, where the depth range of the OMZ is two times greater. The Bay of Bengal patterns may take on added significance as OMZs shoal globally.

Levin, LA, Liu KK, Emeis KC, Breitburg DL, Cloern J, Deutsch C, Giani M, Goffart A, Hofmann EE, Lachkar Z, Limburg K, Liu SM, Montes E, Naqvi W, Ragueneau O, Rabouille C, Sarkar SK, Swaney DP, Wassman P, Wishner KF.  2015.  Comparative biogeochemistry-ecosystem-human interactions on dynamic continental margins. Journal of Marine Systems. 141:3-17.   10.1016/j.jmarsys.2014.04.016   AbstractWebsite

The oceans' continental margins face strong and rapid change, forced by a combination of direct human activity, anthropogenic CO2-induced climate change, and natural variability. Stimulated by discussions in Goa, India at the IMBER IMBIZO III, we (1) provide an overview of the drivers of biogeochemical variation and change on margins, (2) compare temporal trends in hydrographic and biogeochemical data across different margins, (3) review ecosystem responses to these changes, (4) highlight the importance of margin time series for detecting and attributing change and (5) examine societal responses to changing margin biogeochemistry and ecosystems. We synthesize information over a wide range of margin settings in order to identify the commonalities and distinctions among continental margin ecosystems. Key drivers of biogeochemical variation include long-term climate cycles, CO2-induced warming, acidification, and deoxygenation, as well as sea level rise, eutrophication, hydrologic and water cycle alteration, changing land use, fishing, and species invasion. Ecosystem responses are complex and impact major margin services. These include primary production, fisheries production, nutrient cycling, shoreline protection, chemical buffering, and biodiversity. Despite regional differences, the societal consequences of these changes are unarguably large and mandate coherent actions to reduce, mitigate and adapt to multiple stressors on continental margins. (C) 2014 Elsevier BM. All rights reserved.

2014
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 mol.kg(-1)) than those exposed to higher ambient pHT (7.92-7.94) and [O-2] (241-243 mu mol.kg(-1)). In a separate experiment, exposure to low pHT (7.55-7.56) or low [O-2] (83-86 mu mol.kg(-1)) 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.

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

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

Nordstrom, MC, Currin CA, Talley TS, Whitcraft CR, Levin LA.  2014.  Benthic food-web succession in a developing salt marsh. Marine Ecology Progress Series. 500:43-U69.   10.3354/meps10686   AbstractWebsite

Ecological succession has long been a focal point for research, and knowledge of underlying mechanisms is required if scientists and managers are to successfully promote recovery of ecosystem function following disturbance. We addressed the influence of bottom-up processes on successional assemblage shifts in salt marshes, ecosystems with strong physical gradients, and how these shifts were reflected in the trophic characteristics of benthic fauna. We tracked the temporal development of infaunal community structure and food-web interactions in a young, created salt marsh and an adjacent natural marsh in Mission Bay, California, USA (1996-2003). Macro faunal community succession in created Spartina foliosa habitats occurred rapidly, with infaunal densities reaching 70% of those in the natural marsh after 1 yr. Community composition shifted from initial dominance of insect larvae (surface-feeding microalgivores) to increased dominance of oligo chaetes (subsurface-feeding detritivores) within the first 7 yr. Isotopic labeling of microalgae, N-2-fixing cyanobacteria, S. foliosa and bacteria revealed direct links (or absence thereof) between these basal food sources and specific consumer groups. In combination with the compositional changes in the macroinvertebrate fauna, the trophic patterns indicated an increase in food-web complexity over time, reflecting resource-driven marsh succession. Natural abundance stable isotope ratios of salt marsh consumers (infaunal and epifaunal macroinvertebrates, and fish) initially reflected distinctions in trophic structure between the created and natural marsh, but these diminished during successional development. Our findings suggest that changing resource availability is one of the important drivers of succession in benthic communities of restored wetlands in Southern California.

Frieder, CA, Gonzalez JP, Bockmon EE, Navarro MO, Levin LA.  2014.  Can variable pH and low oxygen moderate ocean acidification outcomes for mussel larvae? Global Change Biology. 20:754-764.   10.1111/gcb.12485   AbstractWebsite

Natural variation and changing climate in coastal oceans subject meroplanktonic organisms to broad ranges of pH and oxygen ([O2 ]) levels. In controlled-laboratory experiments we explored the interactive effects of pH, [O2 ], and semidiurnal pH fluctuations on the survivorship, development, and size of early life stages of two mytilid mussels, Mytilus californianus and M. galloprovincialis. Survivorship of larvae was unaffected by low pH, low [O2 ], or semidiurnal fluctuations for both mytilid species. Low pH (<7.6) resulted in delayed transition from the trochophore to veliger stage, but this effect of low pH was absent when incorporating semidiurnal fluctuations in both species. Also at low pH, larval shells were smaller and had greater variance; this effect was absent when semidiurnal fluctuations of 0.3 units were incorporated at low pH for M. galloprovincialis but not for M. californianus. Low [O2 ] in combination with low pH had no effect on larval development and size, indicating that early life stages of mytilid mussels are largely tolerant to a broad range of [O2 ] reflective of their environment (80-260 μmol kg(-1) ). The role of pH variability should be recognized as an important feature in coastal oceans that has the capacity to modulate the effects of ocean acidification on biological responses.

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

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

2013
Neira, C, King I, Mendoza G, Sellanes J, De Ley P, Levin LA.  2013.  Nematode community structure along a central Chile margin transect influenced by the oxygen minimum zone. Deep Sea Research Part I: Oceanographic Research Papers. 78:1-15.   http://dx.doi.org/10.1016/j.dsr.2013.04.002   AbstractWebsite

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

Mora, C, Wei CL, Rollo A, Amaro T, Baco AR, Billett D, Bopp L, Chen Q, Collier M, Danovaro R, Gooday AJ, Grupe BM, Halloran PR, Ingels J, Jones DOB, Levin LA, Nakano H, Norling K, Ramirez-Llodra E, Rex M, Ruhl HA, Smith CR, Sweetman AK, Thurber AR, Tjiputra JF, Usseglio P, Watling L, Wu TW, Yasuhara M.  2013.  Biotic and human vulnerability to projected changes in ocean biogeochemistry over the 21st century. Plos Biology. 11   10.1371/journal.pbio.1001682   AbstractWebsite

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world's ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world's ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.

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

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

2012
Frieder, CA, Nam SH, Martz TR, Levin LA.  2012.  High temporal and spatial variability of dissolved oxygen and pH in a nearshore California kelp forest. Biogeosciences. 9:3917-3930. AbstractWebsite

Predicting consequences of ocean deoxygenation and ocean acidification for nearshore marine ecosystems requires baseline dissolved oxygen (DO) and carbonate chemistry data that are both high-frequency and high-quality. Such data allow accurate assessment of environmental variability and present-day organism exposure regimes. In this study, scales of DO and pH variability were characterized over one year in a nearshore kelp forest ecosystem in the Southern California Bight. DO and pH were strongly, positively correlated, revealing that organisms on this upwelling shelf are not only exposed to low pH but also to low DO. The dominant scale of temporal DO and pH variability occurred on semi-diurnal, diurnal and event (days-weeks) time scales. Daily ranges in DO and pH at 7 m water depth (13 mab) could be as large as 220 mu mol kg(-1) and 0.36 units, respectively. Sources of pH and DO variation include photosynthesis within the kelp forest ecosystem, which can elevate DO and pH by up to 60 mu mol kg(-1) and 0.1 units over one week following the intrusion of high-density, nutrient-rich water. Accordingly, highly productive macrophyte-based ecosystems could serve as deoxygenation and acidification refugia by acting to elevate DO and pH relative to surrounding waters. DO and pH exhibited greater spatial variation over a 10 m increase in water depth (from 7 to 17 m) than along a 5 km stretch of shelf in a cross-shore or alongshore direction. Over a three-month time period, mean DO and pH at 17 m water depth were 168 mu mol kg(-1) and 7.87, respectively. These values represent a 35% decrease in mean DO and 37% increase in [H+] relative to near-surface waters. High-frequency variation was also reduced at depth. The mean daily range in DO and pH was 39% and 37% less, respectively, at 17m water depth relative to 7 m. As a consequence, the exposure history of an organism is largely a function of its depth of occurrence within the kelp forest. With knowledge of local alkalinity conditions and high-frequency temperature, salinity, and pH data, we estimated pCO(2) and calcium carbonate saturation states with respect to calcite and aragonite (Omega(calc) and Omega(arag)) for the La Jolla kelp forest at 7 m and 17 m water depth. pCO(2) ranged from 246 to 1016 mu atm, Omega(calc) was always supersaturated, and Omega(arag) was undersaturated at the beginning of March for five days when pH was less than 7.75 and DO was less than 115 mu mol kg(-1). These findings raise the possibility that the benthic communities along eastern boundary current systems are currently acclimatized and adapted to natural, variable, and low DO and pH. Still, future exposure of coastal California populations to even lower DO and pH may increase as upwelling intensifies and hypoxic boundaries shoal, compressing habitats and challenging the physiological capacity of intolerant species.

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

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

2010
Gooday, AJ, Bett BJ, Escobar E, Ingole B, Levin LA, Neira C, Raman AV, Sellanes J.  2010.  Habitat heterogeneity and its influence on benthic biodiversity in oxygen minimum zones. Marine Ecology-an Evolutionary Perspective. 31:125-147.   10.1111/j.1439-0485.2009.00348.x   AbstractWebsite

Oxygen minimum zones (OMZs; midwater regions with O(2) concentrations <0.5 ml l(-1)) are mid-water features that intercept continental margins at bathyal depths (100-1000 m). They are particularly well developed in the Eastern Pacific Ocean, the Arabian Sea and the Bay of Bengal. Based on analyses of data from these regions, we consider (i) how benthic habitat heterogeneity is manifested within OMZs, (ii) which aspects of this heterogeneity exert the greatest influence on alpha and beta diversity within particular OMZs and (iii) how heterogeneity associated with OMZs influences regional (gamma) diversity on continental margins. Sources of sea-floor habitat heterogeneity within OMZs include bottom-water oxygen and sulphide gradients, substratum characteristics, bacterial mats, and variations in the organic matter content of the sediment and pH. On some margins, hard grounds, formed of phosphorites, carbonates or biotic substrata, represent distinct subhabitats colonized by encrusting faunas. Most of the heterogeneity associated with OMZs, however, is created by strong sea-floor oxygen gradients, reinforced by changes in sediment characteristics and organic matter content. For the Pakistan margin, combining these parameters revealed clear environmental and faunal differences between the OMZ core and the upper and lower boundary regions. In all Pacific and Arabian Sea OMZs examined, oxygen appears to be the master driver of alpha and beta diversity in all benthic faunal groups for which data exist, as well as macrofaunal assemblage composition, particularly in the OMZ core. However, other factors, notably organic matter quantity and quality and sediment characteristics, come into play as oxygen concentrations begin to rise. The influence of OMZs on meiofaunal, macrofaunal and megafaunal regional (gamma) diversity is difficult to assess. Hypoxia is associated with a reduction in species richness in all benthic faunal groups, but there is also evidence for endemism in OMZ settings. We conclude that, on balance, OMZs probably enhance regional diversity, particularly in taxa such as Foraminifera, which are more tolerant of hypoxia than others. Over evolutionary timescales, they may promote speciation by creating strong gradients in selective pressures and barriers to gene flow.