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Grazing impact of the invasive clam Corbula amurensis on the microplankton assemblage of the northern San Francisco Estuary

Grazing by the overbite clam Corbula amurensis (formerly known as Potamocorbula) may be the cause of substantial declines in phytoplankton biomass and zooplankton in the San Francisco Estuary (SFE) following its introduction in 1986. While grazing rates have been examined on bacteria, phytoplankton, and copepod nauplii, the consumption of protistan microzooplankton by C. amurensis has not previously been measured. In this study, laboratory feeding experiments revealed that C. amurensis cleared 0.5 l ind -1 h -1 of microzooplankton (ciliates) and 0.2 l ind -1 h -1 of chlorophyll (chl) a. Despite the higher clearance rate on microzooplankton, clams obtained more of their carbon from phytoplankton, which dominated the prey assemblage on most dates. When the measured clearance rates are extrapolated to field populations of clams, fractional loss rates (50 to 90% d -1 ) exceed the population growth capacity of microzooplankton. Although microzooplankton may not be a major component of the diet of these clams, C. amurensis may further alter food web dynamics through consumption of this important trophic intermediary, thus disrupting this link from bacteria and phytoplankton to higher trophic levels.

California

Marine and terrestrial factors affecting Adélie penguin Pygoscelis adeliae chick growth and recruitment off the western Antarctic Peninsula

An individual-based bioenergetics model that simulates the growth of an Adélie penguin Pygoscelis adeliae chick from hatching to fledging was used to assess marine and terrestrial factors that affect chick growth and fledging mass off the western Antarctic Peninsula. Simulations considered the effects on Adélie penguin fledging mass of (1) modification of chick diet through the addition of Antarctic silverfish Pleuragramma antarcticum to an all-Antarctic krill Euphausia superba diet, (2) reduction of provisioning rate which may occur as a result of an environmental stress such as reduced prey availability, and (3) increased thermoregulatory costs due to wetting of chicks which may result from increased precipitation or snow-melt in colonies. Addition of 17% Antarctic silverfish of Age-Class 3 yr (AC3) to a penguin chick diet composed of Antarctic krill increased chick fledging mass by 5%. Environmental stress that results in >4% reduction in provisioning rate or wetting of just 10% of the chick’s surface area decreased fledging mass enough to reduce the chick’s probability of successful recruitment. The negative effects of reduced provisioning and wetting on chick growth can be compensated for by inclusion of Antarctic silverfish of AC3 and older in the chick diet. Results provide insight into climate-driven processes that influence chick growth and highlight a need for field research designed to investigate factors that determine the availability of AC3 and older Antarctic silverfish to foraging Adélie penguins and the influence of snowfall on chick wetting, thermoregulation and adult provisioning rate.

Marine Ecology Progress Series

Organic carbon sources and sinks in San Francisco Bay: variability induced by river flow

Sources and sinks of organic carbon for San Francisco Bay (California, USA) were estimated for 1980. Sources for the southern reach were dominated by phytoplankton and benthic microalgal production. River loading of organic matter was an additional important factor in the northern reach. Tidal marsh export and point sources played a secondary role. Autochthonous production in San Francisco Bay appears to be less than the mean for temperate-zone estuaries, primarily because turbidity limits microalgal production and the development of seagrass beds. Exchange between the Bay and Pacific Ocean plays an unknown but potentially important role in the organic carbon balance. Interannual variability in the organic carbon supply was assessed for Suisun Bay, a northern reach subembayment that provides habitat for important fish species (delta smelt Hypomesus transpacificus and larval striped bass Morone saxatilus). The total supply fluctuated by an order of magnitude; depending on the year, either autochthonous sources (phytoplankton production) or allochthonous sources (riverine loading) could be dominant. The primary cause of the year-to-year change was variability of freshwater inflows from the Sacramento and San Joaquin rivers, and its magnitude was much larger than long-term changes arising from marsh destruction and point source decreases. Although interannual variability of the total organic carbon supply could not be assessed for the southern reach, year-to-year changes in phytoplankton production were much smaller than in Suisun Bay, reflecting a relative lack of river influence.

California

Long-term effects of the 'Exxon Valdez' oil spill: Sea otter foraging in the intertidal as a pathway of exposure to lingering oil

The protracted recovery of some bird and mammal populations in western Prince William Sound (WPWS), Alaska, and the persistence of spilled 'Exxon Valdez' oil in intertidal sediments, suggests a pathway of exposure to consumers that occupy nearshore habitats. To evaluate the hypothesis that sea otter ( Enhydra lutris ) foraging allows access to lingering oil, we contrast spatial relations between foraging behavior and documented oil distribution. We recovered archival time-depth recorders implanted in 19 sea otters in WPWS, where lingering oil and delayed ecosystem recovery are well documented. Sea otter foraging dives ranged from +2.7 to -92 m below sea level (MLLW), with intertidal accounting for 5 to 38% of all foraging. On average, female sea otters made 16050 intertidal dives per year and 18% of these dives were at depths above the +0.80 m tidal elevation. Males made 4100 intertidal dives per year and 26% of intertidal foraging took place at depths above the +0.80 m tidal elevation. Estimated annual oil encounter rates ranged from 2 to 24 times yr -1 for females, and 2 to 4 times yr -1 for males. Exposure rates increased in spring when intertidal foraging doubled and females were with small pups. In summer 2008, we found sea otter foraging pits on 13.5 of 24.8 km of intertidal shoreline surveyed. Most pits (82%) were within 0.5 m of the zero tidal elevation and 15% were above 0.5 m, the level above which most (65%) lingering oil remains. In August 2008, we detected oil above background concentrations in 18 of 41 (44%) pits excavated by sea otters on beaches with prior evidence of oiling, with total PAH concentrations up to 56000 ng g−1 dry weight. Our estimates of intertidal foraging, the widespread presence of foraging pits in the intertidal, and the presence of oil in and near sea otter foraging pits documents a pathway of exposure from lingering intertidal oil to sea otters foraging in WPWS.

Alaska

Adding the ocean to the study of seabirds: A brief history of at-sea seabird research

We review the history of how research directed towards marine ornithology has led to an appreciation of seabirds as highly specialized marine organisms. Beginning with R. C. Murphy (Pacific), V. C. Wynne-Edwards (Atlantic), and associates in the early 1900s, the research approach grew from an emphasis on seabird single-species ecology to an appreciation of interacting species assemblages and finally to seabirds being considered as important components of marine food webs. After a slow, drawn-out beginning, the initial main impetus for developing the field was a need to map seabird abundance and distribution tied to understanding impacts of continental shelf resource exploitation. Coalescing during the 1970s to 1980s to facilitate this line of research were 6 factors: (1) ability to identify birds at sea; (2) standardization of techniques to quantify abundance; (3) resources and techniques for mapping; (4) appreciation of how scale affects seabird relationships to hydrographic features and patchy prey; (5) development of computing power and appropriate statistics; and (6) seabird biologists becoming embedded in, as well as organizing, multidisciplinary marine research projects. Future advances in understanding the role of seabirds in marine food webs will be made by seabird biologists participating in multidisciplinary projects using grid-like surveys relative to oceanographic features in combination with instrumentation that reveals the finer details of seabird foraging behaviors.

Marine Ecology Progress Series

Variations of transcript profiles between sea otters Enhydra lutris from Prince William Sound, Alaska, and clinically normal reference otters

Development of blood leukocyte gene transcript profiles has the potential to expand condition assessments beyond those currently available to evaluate wildlife health, including sea otters Enhydra lutris , both individually and as populations. The 10 genes targeted in our study represent multiple physiological systems that play a role in immuno-modulation, inflammation, cell protection, tumor suppression, cellular stress-response, xenobiotic metabolizing enzymes, and antioxidant enzymes. These genes can be modified by biological, physical, or anthropogenic impacts and consequently provide information on the general type of stressors present in a given environment. We compared gene transcript profiles of sea otters sampled in 2008 among areas within Prince William Sound impacted to varying degrees by the 1989 ‘Exxon Valdez’ oil spill with those of captive and wild reference sea otters. Profiles of sea otters from Prince William Sound showed elevated transcription in genes associated with tumor formation, cell death, organic exposure, inflammation, and viral exposure when compared to the reference sea otter group, indicating possible recent and chronic exposure to organic contaminants. Sea otters from historically designated oiled areas within Prince William Sound 19 yr after the oil spill had higher transcription of genes associated with tumor formation, cell death, heat shock, and inflammation than those from areas designated as less impacted by the spill.

Alaska

Spatiotemporal associations between Pacific herring spawn and surf scoter spring migration: evaluating a "silver wave" hypothesis

Surf scoters Melanitta perspicillata are sea ducks that aggregate at spawning events of Pacific herring Clupea pallasi and forage on the eggs, which are deposited in abundance during spring at discrete sites. We evaluated whether migrating scoters followed a ‘silver wave’ of resource availability, analogous to the ‘green wave’ of high-quality foraging conditions that herbivorous waterfowl follow during spring migration. We confirmed that herring spawning activity began later in the year at higher latitudes, creating a northward-progressing wave of short-term localized food availability. Using satellite telemetry and aerial surveys, we documented the chronology of scoter spring migration and the use of stopover locations in relation to herring spawn timing and locations. We found that the migration chronology paralleled the northward progression of herring spawning events. Although there was considerable variability in the timing of both scoter migration and the initiation of herring spawning, the processes were related beyond a coincidental northward progression. During migration, 60% of the tracked scoters visited at least 1 spawn site, and those that used spawn sites were located on spawn sites for approximately one-third of their migration locations. Surf scoters showed close spatiotemporal associations with herring spawning events, confirming that the presence of herring spawn was a factor determining habitat use for many individuals. Surf scoters showed close spatiotemporal associations with herring spawning events, confirming that the presence of herring spawn was a factor determining habitat use for many individuals, a conclusion that is consistent with previous studies which used physiologically based metrics to evaluate the importance of herring spawn.

Marine Ecology Progress Series

Bioenergetics model for estimating food requirements of female Pacific walruses ( Odobenus rosmarus divergens )

Pacific walruses Odobenus rosmarus divergens use sea ice as a platform for resting, nursing, and accessing extensive benthic foraging grounds. The extent of summer sea ice in the Chukchi Sea has decreased substantially in recent decades, causing walruses to alter habitat use and activity patterns which could affect their energy requirements. We developed a bioenergetics model to estimate caloric demand of female walruses, accounting for maintenance, growth, activity (active in-water and hauled-out resting), molt, and reproductive costs. Estimates for non-reproductive females 0–12 yr old (65−810 kg) ranged from 16359 to 68960 kcal d −1 (74−257 kcal d −1 kg −1 ) for years with readily available sea ice for which we assumed animals spent 83% of their time in water. This translated into the energy content of 3200–5960 clams per day, equivalent to 7–8% and 14–9% of body mass per day for 5–12 and 2–4 yr olds, respectively. Estimated consumption rates of 12 yr old females were minimally affected by pregnancy, but lactation had a large impact, increasing consumption rates to 15% of body mass per day. Increasing the proportion of time in water to 93%, as might happen if walruses were required to spend more time foraging during ice-free periods, increased daily caloric demand by 6–7% for non-lactating females. We provide the first bioenergetics-based estimates of energy requirements for walruses and a first step towards establishing bioenergetic linkages between demography and prey requirements that can ultimately be used in predicting this population’s response to environmental change.

Marine Ecology Progress Series

Home range, habitat use, and migrations of hawksbill turtles tracked from Dry Tortugas National Park, Florida, USA

To determine habitat-use patterns of sub-adult hawksbills Eretmochelys imbricata , we conducted satellite- and acoustic-tracking of 3 turtles captured in August 2008 within Dry Tortugas National Park (DRTO), south Florida, USA, in the Gulf of Mexico; turtles ranged in size from 51.9 to 69.8 cm straight carapace length. After 263, 699, and 655 d of residence in the park, turtles migrated out of the DRTO. Within the park, core-use areas (i.e. 50% kernel density estimates) were 9.2 to 21.5 km 2 ; all 3 turtle core-use areas overlapped in an area 6.1 km 2 within a zone of the park with multiple human uses (e.g. fishing, anchoring). Two turtles migrated to Cuba and ceased transmitting after 320 and 687 tracking days; the third turtle migrated toward Key West, Florida, and ceased transmitting after 884 tracking days. The present study highlights previously unknown regional connections for hawksbills, possible turtle-harvest incidents, and fine-scale habitat use of sub-adult hawksbills within a United States National Park.

Florida

Walrus areas of use in the Chukchi Sea during sparse sea ice cover

The Pacific walrus Odobenus rosmarus divergens feeds on benthic invertebrates on the continental shelf of the Chukchi and Bering Seas and rests on sea ice between foraging trips. With climate warming, ice-free periods in the Chukchi Sea have increased and are projected to increase further in frequency and duration. We radio-tracked walruses to estimate areas of walrus foraging and occupancy in the Chukchi Sea from June to November of 2008 to 2011, years when sea ice was sparse over the continental shelf in comparison to historical records. The earlier and more extensive sea ice retreat in June to September, and delayed freeze-up of sea ice in October to November, created conditions for walruses to arrive earlier and stay later in the Chukchi Sea than in the past. The lack of sea ice over the continental shelf from September to October caused walruses to forage in nearshore areas instead of offshore areas as in the past. Walruses did not frequent the deep waters of the Arctic Basin when sea ice retreated off the shelf. Walruses foraged in most areas they occupied, and areas of concentrated foraging generally corresponded to regions of high benthic biomass, such as in the northeastern (Hanna Shoal) and southwestern Chukchi Sea. A notable exception was the occurrence of concentrated foraging in a nearshore area of northwestern Alaska that is apparently depauperate in walrus prey. With increasing sea ice loss, it is likely that walruses will increase their use of coastal haul-outs and nearshore foraging areas, with consequences to the population that are yet to be understood.

Alaska;Chukotka

Modelling the seasonality of subsurface light and primary production in the Arabian Sea

Seasonal changes in mixed-layer depth and phytoplankton biomass in the Arabian Sea are assessed with climatologies of ship-based hydrographic measurements and ocean-color observations from satellite. At the close of the intermonsoons in November and especially May, the open Arabian Sea resembles the stereotypic, unperturbed tropical ocean, with a thin oligotrophic mixed layer and a pronounced subsurface chlorophyll maximum. Both the northeast and southwest monsoons disrupt this typical tropical hydrography through mixed-layer deepening and eutrophication in the central and northern Arabian Sea. Computations using a spectral model of light penetration suggest that seasonal changes in mixed-layer thickness and phytoplankton concentration result in pronounced fluctuations through the annual cycle in the radiant flux reaching the base of the mixed layer. At the close of the fall and spring intermonsoons the base of the model euphotic zone is in the thermocline across all of the open Arabian Sea. The euphotic zone appears to rise into the mixed layer of the northern Arabian Sea during both the winter and summer monsoons. Strong seasonality in total primary production and its partitioning between the mixed layer and thermocline is predicted byb a photo-synthesis-irradiance model for a site in the western Arabian Sea (14.36 ° N, 57.38° E). Modeled mixed-layer primary production depicts an intense peak for the southwest monsoon and a secondary northeast monsoon peak separated by intermonsoon period of low production. During the fall and spring intermonsoons, in the presence of a subsurface clorophyll maximum, the model estimate of primary production in the thermocline exceeds that in the mixed layer. Our model calculations suggest that the subsurface clorophyll maximum present in the Arabian Sea during the spring intermonsoon is a precursor of the regional, summer, phytoplankton bloom.

Marine Ecology Progress Series

Mapping spatial resources with GPS animal telemetry: foraging manatees locate seagrass beds in the Ten Thousand Islands, Florida, USA

Turbid water conditions make the delineation and characterization of benthic habitats difficult by traditional in situ and remote sensing methods. Here, we develop and validate modeling and sampling methodology for detecting and characterizing seagrass beds by analyzing GPS telemetry records from radio-tagged manatees. Between October 2002 and October 2005, 14 manatees were tracked in the Ten Thousand Islands (TTI) in southwest Florida (USA) using Global Positioning System (GPS) tags. High density manatee use areas were found to occur off each island facing the open, nearshore waters of the Gulf of Mexico. We implemented a spatially stratified random sampling plan and used a camera-based sampling technique to observe and record bottom observations of seagrass and macroalgae presence and abundance. Five species of seagrass were identified in our study area: Halodule wrightii, Thalassia testudinum, Syringodium filiforme, Halophila engelmannii, and Halophila decipiens. A Bayesian model was developed to choose and parameterize a spatial process function that would describe the observed patterns of seagrass and macroalgae. The seagrasses were found in depths <2 m and in the higher manatee use strata, whereas macroalgae was found at moderate densities at all sampled depths and manatee use strata. The manatee spatial data showed a strong association with seagrass beds, a relationship that increased seagrass sampling efficiency. Our camera-based field sampling proved to be effective for assessing seagrass density and spatial coverage under turbid water conditions, and would be an effective monitoring tool to detect changes in seagrass beds.

Florida

Kittiwake diets and chick production signal a 2008 regime shift in the Northeast Pacific

I examined ~2700 food samples collected from adult and nestling black-legged kittiwakes Rissa tridactyla from 1978 through 2011 on Middleton Island in the Gulf of Alaska. The kittiwake diet was composed chiefly of fish, but invertebrates were taken in appreciable quantities in April and May. Upon spring arrival at the colony, adult kittiwakes foraged regularly at night on vertically migrating mesopelagic prey—lanternfishes (Myctophidae), squids, crustaceans, and polychaetes—a behavior they largely discontinued by egg-laying. During incubation and chick-rearing, food samples contained mostly (~85% by weight) Pacific sand lance Ammodytes hexapterus, capelin Mallotus villosus, Pacific herring Clupea pallasii, sablefish Anopoploma fimbria, krill (Euphausiidae), and juvenile salmon Onchorynchus gorboscha and O. keta. A salient finding over the longitudinal study was the emergence, twice, of capelin as a dominant forage species—once in 2000 to 2003, and again in 2008 through 2011. Kittiwakes responded to capelin availability by producing markedly higher numbers of fledged young. The 2000 to 2003 event corresponded to a previously documented shift to cooler conditions in the NE Pacific, which apparently was relatively limited in magnitude or duration. The more recent transition appears stronger and may be more lasting. I submit that 2008 was an important turning point, marking a substantive reversal of warm conditions that began with the well-documented regime shift of 1977. That interpretation is consistent with the existence of a ~60 yr cycle in ocean and atmospheric conditions in the North Pacific. All else being equal, it predicts the next 20 to 30 yr will be favorable for species such as kittiwakes and Steller sea lions, which seemed to respond negatively to the 1977 to 2007 warm phase of the Pacific Decadal Oscillation.

Alaska

Quantifying long-term risks to sea otters from the 1989 'Exxon Valdez' oil spill: reply to Harwell & Gentile (2013)

Recovery of sea otter populations in Prince William Sound (PWS), Alaska, has been delayed for more than 2 decades following the 1989 ‘Exxon Valdez’ oil spill. Harwell & Gentile (2013; Mar Ecol Prog Ser 488:291–296) question our conclusions in Bodkin et al. (2012; Mar Ecol Prog Ser 447:273-287) regarding adverse effects that oil lingering in the environment may have on sea otters. They agree that exposure may continue, but disagree that it constitutes a significant risk to sea otters. In Bodkin et al. (2012), we suggested that subtle effects of chronic exposure were the most reasonable explanation for delayed recovery of the sea otter population in areas of western PWS, where shorelines were most heavily oiled. Here, we provide additional information on the ecology of sea otters that clarifies why the toxicological effects of oral ingestion of oil do not reflect all effects of chronic exposure. The full range of energetic, behavioral, and toxicological concerns must be considered to appraise how chronic exposure to residual oil may constrain recovery of sea otter populations.

Alaska

Influence of estuarine processes on spatiotemporal variation in bioavailable selenium

Dynamic processes (physical, chemical and biological) challenge our ability to quantify and manage the ecological risk of chemical contaminants in estuarine environments. Selenium (Se) bioavailability (defined by bioaccumulation), stable isotopes and molar carbon-tonitrogen ratios in the benthic clam Potamocorbula amurensis, an important food source for predators, were determined monthly for 17 yr in northern San Francisco Bay. Se concentrations in the clams ranged from a low of 2 to a high of 22 &mu;g g -1 over space and time. Little of that variability was stochastic, however. Statistical analyses and preliminary hydrodynamic modeling showed that a constant mid-estuarine input of Se, which was dispersed up- and down-estuary by tidal currents, explained the general spatial patterns in accumulated Se among stations. Regression of Se bioavailability against river inflows suggested that processes driven by inflows were the primary driver of seasonal variability. River inflow also appeared to explain interannual variability but within the range of Se enrichment established at each station by source inputs. Evaluation of risks from Se contamination in estuaries requires the consideration of spatial and temporal variability on multiple scales and of the processes that drive that variability.

Marine Ecology Progress Series

Estimating movement and survival rates of a small saltwater fish using autonomous antenna receiver arrays and passive integrated transponder tags

We evaluated the performance of small (12.5 mm long) passive integrated transponder (PIT) tags and custom detection antennas for obtaining fine-scale movement and demographic data of mummichog Fundulus heteroclitus in a salt marsh creek. Apparent survival and detection probability were estimated using a Cormack Jolly Seber (CJS) model fitted to detection data collected by an array of 3 vertical antennas from November 2010 to March 2011 and by a single horizontal antenna from April to August 2011. Movement of mummichogs was monitored during the period when the array of vertical antennas was used. Antenna performance was examined in situ using tags placed in wooden dowels (drones) and in live mummichogs. Of the 44 tagged fish, 42 were resighted over the 9 mo monitoring period. The in situ detection probabilities of the drone and live mummichogs were high (~80-100%) when the ambient water depth was less than ~0.8 m. Upstream and downstream movement of mummichogs was related to hourly water depth and direction of tidal current in a way that maximized time periods over which mummichogs utilized the intertidal vegetated marsh. Apparent survival was lower during periods of colder water temperatures in December 2010 and early January 2011 (median estimate of daily apparent survival = 0.979) than during other periods of the study (median estimate of daily apparent survival = 0.992). During late fall and winter, temperature had a positive effect on the CJS detection probability of a tagged mummichog, likely due to greater fish activity over warmer periods. During the spring and summer, this pattern reversed possibly due to mummichogs having reduced activity during the hottest periods. This study demonstrates the utility of PIT tags and continuously operating autonomous detection systems for tracking fish at fine temporal scales, and improving estimates of demographic parameters in salt marsh creeks that are difficult or impractical to sample with active fishing gear.

Marine Ecology Progress Series