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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.

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

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

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.

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.

Neira, C, Delgadillo-Hinojosa F, Zirino A, Mendoza G, Levin LA, Porrachia M, Deheyn DD.  2009.  Spatial distribution of copper in relation to recreational boating in a California shallow-water basin. Chemistry and Ecology. 25:417-433.   10.1080/02757540903334197   AbstractWebsite

The overall effect of the number of boats on the copper (Cu) levels in the water column and sediment, along with their spatial variability within Shelter Island Yacht Basin (SIYB), San Diego Bay, California was examined. We identified a horizontal gradient of increasing dissolved Cu and Cu in sediment from outside to the head of SIYB which was coincident with the increasing number of boats. Spatial models of Cu distribution in water and sediment indicated the presence of 'hotspots' of Cu concentration. From outside to the head of SIYB, dissolved Cu ranged from 1.3 gL-1 to 14.6 gL-1 in surface water, and 2.0 gL-1 to 10.2 gL-1 in bottom water. Cu in sediment exceeded the Effect Range Low of 34mgkg-1 (i.e. where adverse effects to fauna may occur), with a peak concentration of 442mgkg-1 at the head of the basin. Free Cu++ in surface water was several orders of magnitude higher than in sediment porewater. High-resolution data of Cu species together with probability maps presented in this paper will allow managers to easily visualise and localise areas of impaired quality and to prioritise which areas should be targeted to improve Cu-related conditions.

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.

Neira, C, Levin LA, Grosholz ED.  2005.  Benthic macrofaunal communities of three sites in San Francisco Bay invaded by hybrid Spartina, with comparison to uninvaded habitats. Marine Ecology-Progress Series. 292:111-126.   10.3354/meps292111   AbstractWebsite

A hybrid cordgrass, formed from a cross between Spartina alterniflora (Atlantic cordgrass) and S. foliosa (Pacific cordgrass), has recently spread within the intertidal zone of south San Francisco Bay. Sediment properties and macroinfaunal community structure were compared in patches invaded by Spartina hybrid and adjacent uninvaded patches at 3 sites in San Francisco Bay (2 tidal flats and 1 Salicornia marsh). We hypothesized that (1) sediments vegetated by Spartina hybrid would have reduced sediment grain size, higher organic matter content, lower redox potential, lower salinity and reduced microalgal biomass relative to adjacent unvegetated tidal flat sediments, and (2) that differences in the sediment environment would correspond to changes in the infaunal invertebrate community structure and feeding modes. We observed 75 % lower total macro-faunal density and lower species richness in Spartina-vegetated sediments at Elsie Roemer (30 yr old invasion) than in an adjacent unvegetated tidal flat. This was due to lower densities of surface-feeding amphipods, bivalves, cirratulid and spionid polychaetes. The proportional representation of subsurface-deposit feeders was greater in Spartina patches than in unvegetated sediments. At a more recently invaded site (Roberts Landing; 15 yr invasion), Spartina patches differed from tidal flat sediments in composition, but not in abundance. Native (Salicornia) and Spartina patches exhibited similar sediment properties at San Mateo, where the Spartina hybrid invaded 8 to 10 yr earlier. No differences were detected in densities or proportions of surface- or subsurface-deposit feeders, but the proportion of carnivores/omnivores and grazers increased in the hybrid-invaded patches. These studies suggest that the invasive Spartina hybrid in south San Francisco Bay can have differing effects on sediment ecosystems, possibly depending on the location, age, or type of habitats involved.

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.

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.

Neira, C, Grosholz ED, Levin LA, Blake R.  2006.  Mechanisms generating modification of benthos following tidal flat invasion by a Spartina hybrid. Ecological Applications. 16:1391-1404.   10.1890/1051-0761(2006)016[1391:mgmobf];2   AbstractWebsite

Many coastal habitats are being substantially altered by introduced plants. In San Francisco Bay,, California, USA, a hybrid form of the eastern cordgrass Spartina alterniflora is rapidly invading open mudflats in southern and central sections of the Bay, altering habitat, reducing macrofaunal densities, and shifting species composition. The invasion has resulted in significant losses of surface-feeding amphipods, bivalves, and cirratulid polychaetes, while subsurface feeding groups such as tubificid oligochaetes and capitellid polychaetes have been unaffected. In the present paper, we document the causes and mechanisms underlying the changes observed. Through a series of in situ manipulative experiments we examined the influence of hybrid Spartina canopy on a range of physical, chemical, and biological properties. The hybrid Spartina canopy exerted a strong influence on. the hydrodynamic regime, triggering a series of physical, chemical, and biological changes in the benthic system. Relative to tidal flats, water velocity was reduced in hybrid patches, promoting deposition of fine-grained, organic-rich particles. The resulting changes in the sediment environment included increased porewater sulfide concentrations and anoxia, which led to poor survivorship. of surface feeders such as, bivalves, amphipods, and polychaetes. These are key taxa that support higher trophic levels including migratory shorebirds that feed on tidal flats. Altered flow in the Spartina canopy further contributed to changes in barnacle recruitment and resuspension of adult benthic invertebrates. Increased crab-induced predation pressure associated with Spartina invasion also contributed to changes in benthic invertebrate communities. Our results suggest that multiple physical, chemical, biotic, and trophic impacts of the Spartina invasion have resulted in substantial changes in benthic communities that are likely to have important effects on the entire ecosystem.

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

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

Neira, C, Levin LA, Grosholz ED, Mendoza G.  2007.  Influence of invasive Spartina growth stages on associated macrofaunal communities. Biological Invasions. 9:975-993.   10.1007/s10530-007-9097-x   AbstractWebsite

In coastal wetlands, invasive plants often act as ecosystem engineers altering flow, light and sediments which, in turn, can affect benthic animal communities. However, the degree of influence of the engineer will vary significantly as it grows, matures and senesces, and surprisingly little is known about how the influence of an ecosystem engineer varies with ontogeny. We address this issue on the tidal flats of San Francisco Bay where hybrid Spartina (foliosa x alterniflora) invaded 30 years ago. The invasion has altered the physico-chemical properties of the sediment habitat, which we predicted should cause changes in macrofaunal community structure and function. Through mensurative and manipulative approaches we investigated the influence of different growth stages of hybrid Spartina on macrobenthos and the underlying mechanisms. Cross-elevation sampling transects were established covering 5 zones (or stages) of the invasion, running from the tidal flat (pre-invasion) to an unvegetated dieback zone. Additionally, we experimentally removed aboveground plant structure in the mature (inner) marsh to mimic the 'unvegetated areas'. Our results revealed four distinct faunal assemblages, which reflected Spartina-induced changes in the corresponding habitat properties along an elevation gradient: a pre-invaded tidal flat, a leading edge of immature invasion, a center of mature invasion, and a senescing dieback area. These stages of hybrid Spartina invasion were accompanied by a substantial reduction in macrofaunal species richness and an increase in dominance, as well as a strong shift in feeding modes, from surface microalgal feeders to subsurface detritus/Spartina feeders (mainly tubificid oligochaetes and capitellid polychaetes). Knowledge of the varying influence of plant invaders on the sediment ecosystem during different phases of invasion is critical for management of coastal wetlands.

Neira, C, Sellanes J, Levin LA, Arntz WE.  2001.  Meiofaunal distributions on the Peru margin: relationship to oxygen and organic matter availability. Deep-Sea Research Part I-Oceanographic Research Papers. 48:2453-2472.   10.1016/s0967-0637(01)00018-8   AbstractWebsite

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12 degreesS). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440-548 ind. 10cm(-2) (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2-5 cm. interval. At the deeper sites, about 50% were restricted to the top I cm. The importance of copepods and nauplii increased consistently with depth, reaching similar to 12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02mll(-1) supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r = 0.99, p <0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic. heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites. (C) 2001 Elsevier Science Ltd. All rights reserved.

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.

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.

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.

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.

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.