Showing papers in "Deep-sea Research Part Ii-topical Studies in Oceanography in 2014"

Megafauna of vulnerable marine ecosystems in French mediterranean submarine canyons: Spatial distribution and anthropogenic impacts

Abstract: Vulnerable Marine Ecosystems (VME) in the deep Mediterranean Sea have been identified by the General Fisheries Commission for the Mediterranean as consisting of communities of Scleractinia (Lophelia pertusa and Madrepora oculata), Pennatulacea (Funiculina quadrangularis) and Alcyonacea (Isidella elongata). This paper deals with video data recorded in the heads of French Mediterranean canyons. Quantitative observations were extracted from 101 video films recorded during the MEDSEACAN cruise in 2009 (Aamp/Comex). Qualitative information was extracted from four other cruises (two Marum/Comex cruises in 2009 and 2011 and two Ifremer cruises in 1995 and 2010) to support the previous observations in the Cassidaigne and Lacaze-Duthiers canyons. All the species, fishing impacts and litter recognized in the video films recorded from 180 to 700 m depth were mapped using GIS. The abundances and distributions of benthic fishing resources (marketable fishes, Aristeidae, Octopodidae), Vulnerable Marine Species, trawling scars and litter of 17 canyons were calculated and compared, as was the open slope between the Stoechades and Toulon canyons. Funiculina quadrangularis was rarely observed, being confined for the most part to the Marti canyon and, I. elongata was abundant in three canyons (Bourcart, Marti, Petit-Rhone). These two cnidarians were encountered in relatively low abundances, and it may be that they have been swept away by repeated trawling. The Lacaze-Duthiers and Cassidaigne canyons comprised the highest densities and largest colony sizes of scleractinian cold-water corals, whose distribution was mapped in detail. These colonies were often seen to be entangled in fishing lines. The alcyonacean Callogorgia verticillata was observed to be highly abundant in the Bourcart canyon and less abundant in several other canyons. This alcyonacean was also severely affected by bottom fishing gears and is proposed as a Vulnerable Marine Species. Our studies on anthropogenic impacts show that seafloor disturbance by benthic fishing is mainly attributable to trawling in the Gulf of Lion and to long lines where rocky substrates are present. The bauxite residue (red mud) expelled in the Cassidaigne canyon was seen to prevent fauna from settling at the bottom of the canyon and it covered much of the flanks. Litter was present in all of the canyons and especially in considerable quantities in the Ligurian Sea, where the heads of the canyons are closer to the coast. Three Marine Protected Areas and one fishing area with restricted access have recently been established and should permit the preservation of these deep ecosystems.

Phytoplankton blooms beneath the sea ice in the Chukchi sea

Abstract: In the Arctic Ocean, phytoplankton blooms on continental shelves are often limited by light availability, and are therefore thought to be restricted to waters free of sea ice. During July 2011 in the Chukchi Sea, a large phytoplankton bloom was observed beneath fully consolidated pack ice and extended from the ice edge to 4100 km into the pack. The bloom was composed primarily of diatoms, with biomass reaching 1291 mg chlorophyll a m � 2 and rates of carbon fixation as high as 3.7 g C m � 2 d � 1 . Although the sea ice where the bloom was observed was near 100% concentration and 0.8–1.2 m thick, 30–40% of its surface was covered by melt ponds that transmitted 4-fold more light than adjacent areas of bare ice, providing sufficient light for phytoplankton to bloom. Phytoplankton growth rates associated with the under-ice bloom averaged 0.9 d � 1

Eddy properties in the Mozambique Channel: A comparison between observations and two numerical ocean circulation models

Abstract: Analysis of satellite altimetry observations, transports estimates from a mooring array, as well as output from two different numerical ocean circulation models (ROMS and HYCOM), have been used to investigate the mesoscale eddy properties and transport variability in the Mozambique Channel. The power spectral density of model transports at 17°S indicates the models ability to represent the transport variability at mesoscale frequencies (range between 3 yr −1 and 10 yr −1 ). The models have shown an exaggerated representation of the lower frequencies (~ −1 ), while underestimating the higher frequency signals (~ >10 yr −1 ). The overestimation of the seasonal cycle appears in our case not to be related to a misrepresentation of the mesoscale variability. The eddies were identified using an automatic eddy tracking scheme. Both anticyclonic and cyclonic eddies appeared to have a preferred site of formation within the channel. The density distribution showed that the anticyclones exhibited a bi-modal distribution: the first mode was associated with the typical scale for the oceanic mesoscale turbulence, while the second mode was related to the passage of large rings at a frequency of about 4–7 per year. On the other hand, cyclonic eddies had a single mode distribution that follows the first baroclinic Rossby radius of deformation, which is a typical scale for the oceanic mesoscale surface eddy variability, suggesting that their formation is associated with baroclinic instability. Eddy mean amplitudes per class of radius (

The geochemistry of deep-sea coral skeletons: A review of vital effects and applications for palaeoceanography

Abstract: Deep-sea corals were discovered over a century ago, but it is only over recent years that focused efforts have been made to explore the history of the oceans using the geochemistry of their skeletal remains. They offer a promising archive of past oceanic environments given their global distribution, layered growth patterns, longevity and preservation as well as our ability to date them using radiometric techniques. This paper provides an overview of the current state-of-the-art in terms of geochemical approaches to using deep-sea coral skeletons to explore the history of the ocean. Deep-sea coral skeletons have a wide array of morphologies (e.g. solitary cup corals, branching colonial corals) and materials (calcite, aragonite and proteins). As such their biomineralization strategies are diverse, leading to complex geochemistry within coral skeletons. Notwithstanding these complications, progress has been made on developing methods for reconstructing the oceanographic environment in the past using trace elements and isotopic methods. Promising approaches within certain coral groups include clumped isotopes and Mg/Li for temperature reconstructions, boron isotopes and radiocarbon for carbon cycling, eNd, and radiocarbon for circulation studies and δ^(15)N, P/Ca and Ba/Ca for nutrient tracer studies. Likewise there is now a range of techniques for dating deep-sea corals skeletons (e.g. U-series, radiocarbon), and determining their growth rates (e.g. radiocarbon and ^(210)Pb). Dating studies on historic coral populations in the Atlantic, Southern Ocean and Pacific point to climate and environmental changes being dominant controls on coral populations over millennial and orbital timescales. This paper provides a review of a range of successes and promising approaches. It also highlights areas in which further research would likely provide new insights into biomineralization, palaeoceanography and distribution of past coral populations.

Alexandrium fundyense cysts in the Gulf of Maine: long-term time series of abundance and distribution, and linkages to past and future blooms.

Abstract: Here we document Alexandrium fundyense cyst abundance and distribution patterns over nine years (1997 and 2004-2011) in the coastal waters of the Gulf of Maine (GOM) and identify linkages between those patterns and several metrics of the severity or magnitude of blooms occurring before and after each autumn cyst survey. We also explore the relative utility of two measures of cyst abundance and demonstrate that GOM cyst counts can be normalized to sediment volume, revealing meaningful patterns equivalent to those determined with dry weight normalization. Cyst concentrations were highly variable spatially. Two distinct seedbeds (defined here as accumulation zones with > 300 cysts cm-3) are evident, one in the Bay of Fundy (BOF) and one in mid-coast Maine. Overall, seedbed locations remained relatively constant through time, but their area varied 3-4 fold, and total cyst abundance more than 10 fold among years. A major expansion of the mid-coast Maine seedbed occurred in 2009 following an unusually intense A. fundyense bloom with visible red-water conditions, but that feature disappeared by late 2010. The regional system thus has only two seedbeds with the bathymetry, sediment characteristics, currents, biology, and environmental conditions necessary to persist for decades or longer. Strong positive correlations were confirmed between the abundance of cysts in both the 0-1 and the 0-3 cm layers of sediments in autumn and geographic measures of the extent of the bloom that occurred the next year (i.e., cysts → blooms), such as the length of coastline closed due to shellfish toxicity or the southernmost latitude of shellfish closures. In general, these metrics of bloom geographic extent did not correlate with the number of cysts in sediments following the blooms (blooms → cysts). There are, however, significant positive correlations between 0-3 cm cyst abundances and metrics of the preceding bloom that are indicative of bloom intensity or vegetative cell abundance (e.g., cumulative shellfish toxicity, duration of detectable toxicity in shellfish, and bloom termination date). These data suggest that it may be possible to use cyst abundance to empirically forecast the geographic extent of the forthcoming bloom and, conversely, to use other metrics from bloom and toxicity events to forecast the size of the subsequent cyst population as the inoculum for the next year's bloom. This is an important step towards understanding the excystment/encystment cycle in A. fundyense bloom dynamics while also augmenting our predictive capability for this HAB-forming species in the GOM.

Trawling-induced daily sediment resuspension in the flank of a Mediterranean submarine canyon

Abstract: Commercial bottom trawling is one of the anthropogenic activities causing the biggest impact on the seafloor due to its recurrence and global distribution. In particular, trawling has been proposed as a major driver of sediment dynamics at depths below the reach of storm waves, but the issue is at present poorly documented with direct observations. This paper analyses changes in water turbidity in a tributary valley of the La Fonera (=Palamos) submarine canyon, whose flanks are routinely exploited by a local trawling fleet down to depths of 800 m. A string of turbidimeters was deployed at 980 m water depth inside the tributary for two consecutive years, 2010–2011. The second year, an ADCP profiled the currents 80 m above the seafloor. The results illustrate that near-bottom water turbidity at the study site is heavily dominated, both in its magnitude and temporal patterns, by trawling-induced sediment resuspension at the fishing ground. Resuspended sediments are channelised along the tributary in the form of sediment gravity flows, being recorded only during working days and working hours of the trawling fleet. These sediment gravity flows generate turbid plumes that extend to at least 100 m above the bottom, reaching suspended sediment concentrations up to 236 mg l−1 close to the seafloor (5 m above bottom). A few hours after the end of daily trawling activities, water turbidity progressively decreases but resuspended particles remain in suspension for several hours, developing bottom and intermediate nepheloid layers that reach background levels ∼2 mg l−1 before trawling activities resume. The presence of these nepheloid layers was recorded in a CTD+turbidimeter transect conducted across the fishing ground a few hours after the end of a working day. These results highlight that deep bottom trawling can effectively replace natural processes as the main driving force of sediment resuspension on continental slope regions and generate increased near-bottom water turbidity that propagates from fishing grounds to wider and deeper areas via sediment gravity flows and nepheloid layer development.

Climate-mediated changes in zooplankton community structure for the eastern Bering Sea

Abstract: Zooplankton are critical to energy transfer between higher and lower trophic levels in the eastern Bering Sea ecosystem. Previous studies from the southeastern Bering Sea shelf documented substantial differences in zooplankton taxa in the Middle and Inner Shelf Domains between warm and cold years. Our investigation expands this analysis into the northern Bering Sea and the south Outer Domain, looking at zooplankton community structure during a period of climate-mediated, large-scale change. Elevated air temperatures in the early 2000s resulted in regional warming and low sea-ice extent in the southern shelf whereas the late 2000s were characterized by cold winters, extensive spring sea ice, and a well-developed pool of cold water over the entire Middle Domain. The abundance of large zooplankton taxa such as Calanus spp. ( C. marshallae and C. glacialis) , and Parasagitta elegans , increased from warm to cold periods, while the abundance of gelatinous zooplankton (Cnidaria) and small taxa decreased. Biomass followed the same trends as abundance, except that the biomass of small taxa in the southeastern Bering Sea remained constant due to changes in abundance of small copepod taxa (increases in Acartia spp. and Pseudocalanus spp. and decreases in Oithona spp.). Statistically significant changes in zooplankton community structure and individual species were greatest in the Middle Domain, but were evident in all shelf domains, and in both the northern and southern portions of the eastern shelf. Changes in community structure did not occur abruptly during the transition from warm to cold, but seemed to begin gradually and build as the influence of the sea ice and cold water temperatures persisted. The change occurred one year earlier in the northern than the southern Middle Shelf. These and previous observations demonstrate that lower trophic levels within the eastern Bering Sea respond to climate-mediated changes on a variety of time scales, including those shorter than the commonly accepted quasi-decadal time periods. This lack of resilience or inertia at the lowest trophic levels affects production at higher trophic levels and must be considered in management strategy evaluations of living marine resources.

Spring and fall phytoplankton blooms in a productive subarctic ecosystem, the eastern Bering Sea, during 1995–2011

Abstract: The timing and magnitude of phytoplankton blooms in subarctic ecosystems often strongly influence the amount of energy that is transferred through subsequent trophic pathways. In the eastern Bering Sea, spring bloom timing has been linked to ice retreat timing and production of zooplankton and fish. A large part of the eastern Bering Sea shelf (~500 km wide) is ice-covered during winter and spring. Four oceanographic moorings have been deployed along the 70-m depth contour of the eastern Bering Sea shelf with the southern location occupied annually since 1995, the two northern locations since 2004 and the remaining location since 2001. Chlorophyll a fluorescence data from the four moorings provide 37 realizations of a spring bloom and 33 realizations of a fall bloom. We found that in the eastern Bering Sea: if ice was present after mid-March, spring bloom timing was related to ice retreat timing ( p n =11). A fall bloom also commonly occurred, usually in late September (average day 274, SE=4.2, n =33), and its timing was not significantly related to the timing of storms ( p =0.88, df=1, 27) or fall water column overturn ( p =0.49, df=1, 27). The magnitudes of the spring and fall blooms were correlated ( p =0.011, df=28). The interval between the spring and fall blooms varied between four to six months depending on year and location. We present a hypothesis to explain how the large crustacean zooplankton taxa Calanus spp. likely respond to variation in the interval between blooms (spring to fall and fall to spring).

Spatial scale-dependent habitat heterogeneity influences submarine canyon macrofaunal abundance and diversity off the Main and Northwest Hawaiian Islands

Abstract: The mapping of biodiversity on continental margins on landscape scales is highly relevant to marine spatial planning and conservation. Submarine canyons are widespread topographic features on continental and island margins that enhance benthic biomass across a range of oceanic provinces and productivity regimes. However, it remains unclear whether canyons enhance faunal biodiversity on landscape scales relevant to marine protected area (MPA) design. Furthermore, it is not known which physical attributes and heterogeneity metrics can provide good surrogates for large-scale mapping of canyon benthic biodiversity. To test mechanistic hypotheses evaluating the role of different canyon-landscape attributes in enhancing benthic biodiversity at different spatial scales we conducted 34 submersible dives in six submarine canyons and nearby slopes in the Hawaiian archipelago, sampling infaunal macrobenthos in a depth-stratified sampling design. We employed multivariate multiple regression models to evaluate sediment and topographic heterogeneity, canyon transverse profiles, and overall water mass variability as potential drivers of macrobenthic community structure and species richness. We find that variables related to habitat heterogeneity at medium (0.13 km2) and large (15–33 km2) spatial scales such as slope, backscatter reflectivity and canyon transverse profiles are often good predictors of macrobenthic biodiversity, explaining 16–30% of the variance. Particulate organic carbon (POC) flux and distance from shore are also important variables, implicating food supply as a major predictor of canyon biodiversity. Canyons off the high Main Hawaiian Islands (Oahu and Moloka'i) are significantly affected by organic enrichment, showing enhanced infaunal macrobenthos abundance, whereas this effect is imperceptible around the low Northwest Hawaiian Islands (Nihoa and Maro Reef). Variable canyon alpha-diversity and high rates of species turnover (beta-diversity), particularly for polychaetes, suggest that canyons play important roles in maintaining high levels of regional biodiversity in the extremely oligotrophic system of the North Pacific Subtropical Gyre. This information is of key importance to the process of MPA design, suggesting that canyon habitats be explicitly included in marine spatial planning.

Submarine canyons as important habitat for cetaceans, with special reference to the Gully: A review

Abstract: There has been much research interest in the use of submarine canyons by cetaceans, particularly beaked whales (family Ziphiidae), which appear to be especially attracted to canyon habitats in some areas. However, not all submarine canyons are associated with large numbers of cetaceans and the mechanisms through which submarine canyons may attract cetaceans are not clearly understood. This paper reviews some of the cetacean associations with submarine canyons that have been anecdotally described or presented in scientific literature and discusses the physical, oceanographic and biological mechanisms that may lead to enhanced cetacean abundance around these canyons. Particular attention is paid to the Gully, a large submarine canyon and Marine Protected Area off eastern Canada for which there exists some of the strongest evidence available for submarine canyons as important cetacean habitat. Studies demonstrating increased cetacean abundance in the Gully and the processes that are likely to attract cetaceans to this relatively well-studied canyon are discussed. This review provides some limited evidence that cetaceans are more likely to associate with larger canyons; however, further studies are needed to fully understand the relationship between the physical characteristics of canyons and enhanced cetacean abundance. In general, toothed whales (especially beaked whales and sperm whales) appear to exhibit the strongest associations with submarine canyons, occurring in these features throughout the year and likely attracted by concentrating and aggregating processes. By contrast, baleen whales tend to occur in canyons seasonally and are most likely attracted to canyons by enrichment and concentrating processes. Existing evidence thus suggests that at least some submarine canyons are important foraging areas for cetaceans, and should be given special consideration for cetacean conservation and protection.

Short-term metabolic and growth responses of the cold-water coral Lophelia pertusa to ocean acidification

Abstract: Cold-water corals are associated with high local biodiversity, but despite their importance as ecosystem engineers, little is known about how these organisms will respond to projected ocean acidification. Since preindustrial times, average ocean pH has decreased from 8.2 to ~8.1, and predicted CO2 emissions will decrease by up to another 0.3 pH units by the end of the century. This decrease in pH may have a wide range of impacts upon marine life, and in particular upon calcifiers such as cold-water corals. Lophelia pertusa is the most widespread cold-water coral (CWC) species, frequently found in the North Atlantic. Here, we present the first short-term (21 days) data on the effects of increased CO2 (750 ppm) upon the metabolism of freshly collected L. pertusa from Mingulay Reef Complex, Scotland, for comparison with net calcification. Over 21 days, corals exposed to increased CO2 conditions had significantly lower respiration rates (11.4±1.39 SE, µmol O2 g−1 tissue dry weight h−1) than corals in control conditions (28.6±7.30 SE µmol O2 g−1 tissue dry weight h−1). There was no corresponding change in calcification rates between treatments, measured using the alkalinity anomaly technique and 14C uptake. The decrease in respiration rate and maintenance of calcification rate indicates an energetic imbalance, likely facilitated by utilisation of lipid reserves. These data from freshly collected L. pertusa from the Mingulay Reef Complex will help define the impact of ocean acidification upon the growth, physiology and structural integrity of this key reef framework forming species.

Predicting the distribution of vulnerable marine ecosystems in the deep sea using presence-background models

Abstract: In 2006 the United Nations called on states to implement measures to prevent significant adverse impacts to vulnerable marine ecosystems (VMEs) in the deep sea. It has been widely recognised that a major limitation to the effective application of these measures to date is uncertainty regarding the distribution of VMEs. Conservationists, researchers, resource managers and governmental bodies are increasingly turning to predictive species distribution models (SDMs) to identify the potential presence of species in areas that have not been sampled. In particular, the development of robust ‘presence-background’ model algorithms has accelerated the application of these techniques for working with presence-only species data. This has allowed scientists to exploit the large amounts of species data held in global biogeographic databases. Despite improvements in model algorithms, environmental data and species presences, there are still limitations to the reliability of these techniques, especially in poorly studied areas such as the deep sea. Recent studies have begun to address a key limitation, the quality of data, by using multibeam echosounder surveys and species data from video surveys to acquire high-resolution data. Whilst these data are often amongst the very best that can be acquired, the surveys are highly localised, often targeted towards known VME-containing areas, are very expensive and time consuming. It is financially prohibitive to survey whole regions or ocean basins using these techniques, so alternative cost-effective approaches are required. Here, we review ‘presence-background’ SDMs in the context of those studies conducted in the deep sea. The issues of sampling bias, spatial autocorrelation, spatial scale, model evaluation and validation are considered in detail, and reference is made to recent developments in species distribution modelling literature. Further information is provided on how these approaches are being used to influence ocean management, and best practises are outlined to aid the effective adoption of these techniques in the future.

Species-specific physiological response by the cold-water corals Lophelia pertusa and Madrepora oculata to variations within their natural temperature range

Abstract: The scleractinian cold-water corals (CWC) Lophelia pertusa and Madrepora oculata represent two major deep-sea reef-forming species that act as key ecosystem engineers over a wide temperature range, extending from the northern Atlantic (ca. 5–9 °C) to the Mediterranean Sea (ca. 11–13 °C). Recent research suggests that environmental parameters, such as food supply, settling substrate availability or aragonite saturation state may represent important precursors controlling habitat suitability for CWC. However, the effect of one principal environmental factor, temperature, on CWC key physiological processes is still unknown. In order to evaluate this effect on calcification, respiration, and dissolved organic carbon (DOC) net flux, colonies of Mediterranean L. pertusa and M. oculata were acclimated in aquaria to three temperatures (12, 9 and 6 °C), by consecutive decrements of 1 month duration. L. pertusa and M. oculata maintained at Mediterranean control conditions (i.e. 12 °C) displayed constant rates, on average respiring 4.8 and 4.0 µmol O2 cm−2 coral surface area d−1, calcifying 22.3 and 12.3 µmol CaCO3 g−1 skeletal dry weight d−1 and net releasing 2.6 and 3.1 µmol DOC cm−2 coral surface area d−1, respectively. Respiration of L. pertusa was not affected by lowered temperatures, while M. oculata respiration declined significantly (by 48%) when temperature decreased to 9 °C and 6 °C relative to controls. L. pertusa calcification at 9 °C was similar to controls, but decreased significantly (by 58%) at 6 °C. For M. oculata, calcification declined by 41% at 9 °C and by 69% at 6 °C. DOC net flux was similar throughout the experiment for both CWC. These findings reveal species-specific physiological responses by CWC within their natural temperature range. L. pertusa shows thermal acclimation in respiration and calcification, while these mechanisms appear largely absent in M. oculata. Conclusively, species-specific thermal acclimation may significantly affect the occurrence and local abundance of cosmopolitan CWC species, consequently influencing their important role in habitat engineering and ecosystem functioning in various thermal environments.

Geochemical and physical constraints for the occurrence of living cold-water corals

Abstract: Cold-water coral communities cover a wide range of possible habitats in terms of latitude, ocean basins, and depth, with ongoing studies continually expanding occurrences in various regions of the global ocean. A range of factors determines the formation of cold-water coral reefs, such as physical, hydrochemical, and biological (e.g. food supply) factors. Recently, more and more modeling studies have emerged using a variety of mathematical approaches have emerged including environmental niche factor analysis (ENFA) and predictive habitat suitability models. However, only few studies have attempted to characterize the underlying suite of hydro-biogeochemical and physical constraints of cold-water coral reefs and to differentiate between pristine reef growth vs. sites with reduced or no coral occurrences. This study concentrates on new data and a compilation of existing data sets on the physical and chemical properties in the NE Atlantic and the Mediterranean. It explores the influence of ambient bottom waters and its characteristics on living cold-water reefs and mounds formed by Lophelia pertusa. Several questions are addressed: (1) what are the physical and geochemical boundary conditions of living cold-water corals? (2) Do these geochemical parameters correlate with proposed physical prerequisites? (3) Is there a general difference in the signature of living and dead coral sites?

The West Melilla cold water coral mounds, Eastern Alboran Sea: Morphological characterization and environmental context

Abstract: A new mound field, the West Melilla mounds, interpreted as being cold-water coral mounds, has been recently unveiled along the upper slope of the Mediterranean Moroccan continental margin, a few kilometers west of the Cape Tres Forcas. This study is based on the integration of high-resolution geophysical data (swath bathymetry, parametric sub-bottom profiler), CTD casts, Acoustic Doppler Current Profiler (ADCP), ROV video and seafloor sampling, acquired during the TOPOMED GASSIS (2011) and MELCOR (2012) cruises. Up to 103 mounds organized in two main clusters have been recognized in a depth range of 299–590 m, displaying a high density of 5 mounds/km2. Mounds, 1–48 m high above the surrounding seafloor and on average 260 m wide, are actually buried by a 1–12 m thick fine-grained sediment blanket. Seismic data suggest that the West Melilla mounds grew throughout the Early Pleistocene–Holocene, settling on erosive unconformities and mass movement deposits. During the last glacial–interglacial transition, the West Melilla mounds may have suffered a drastic change of the local sedimentary regime during the late Holocene and, unable to stand increasing depositional rates, were progressively buried. At the present day, temperature and salinity values on the West Melilla mounds suggest a plausible oceanographic setting, suitable for live CWCs. Nonetheless, more data is required to groundtruth the West Melilla mounds and better constrain the interplay of sedimentary and oceanographic factors during the evolution of the West Melilla mounds.

Evidence of under-ice phytoplankton blooms in the Chukchi Sea from 1998 to 2012

Abstract: The discovery in 2011 of a massive phytoplankton bloom underneath first-year sea ice in the western Arctic has prompted an investigation of the spatial and temporal distribution of under-ice phytoplankton blooms. Here, we explore the satellite record from years 1998 to 2012 for evidence of under-ice blooms on the Chukchi Sea shelf. Phytoplankton blooms were categorized as under-ice blooms, probable under-ice blooms, or marginal ice zone blooms, depending on bloom timing in relation to the timing of ice retreat. Annual bloom type maps reveal that under-ice phytoplankton blooms were present in every year of the satellite record. Averaged over all years, the combination of under-ice blooms and probable under-ice blooms covered a portion of the observable study area that was 2.5-fold higher than that of marginal ice zone blooms (71.5% and 28.5%, respectively). This finding strongly contradicts the traditional view that phytoplankton in seasonally ice-covered waters bloom only after ice retreat and instead indicates that blooms are initiated whenever light and nutrient availability is sufficient for photosynthesis, a condition often reached early in the season underneath first-year sea ice on nutrient-rich continental shelves. Spatial patterns in bloom type were distinguished relative to the date of ice retreat, with probable under-ice blooms dominating the nutrient-rich western Chukchi Sea and at higher latitudes where ice retreats later, while marginal ice zone blooms were most common in the southern and eastern Chukchi Sea where ice retreats earlier. Our results suggest that under-ice phytoplankton blooms are widespread in the Chukchi Sea and had been prevalent there for more than a decade prior to their discovery in 2011.

Diversity and toxicity of the diatom Pseudo-nitzschia Peragallo in the Gulf of Maine, Northwestern Atlantic Ocean

Abstract: Multiple species in the toxic marine diatom genus Pseudo-nitzschia have been identified in the Northwestern Atlantic region encompassing the Gulf of Maine (GOM), including the Bay of Fundy (BOF). To gain further knowledge of the taxonomic composition and toxicity of species in this region, Pseudo-nitzschia isolates (n=146) were cultured from samples collected during research cruises that provided broad spatial coverage across the GOM and the southern New England shelf, herein referred to as the GOM region, during 2007–2008. Isolates, and cells in field material collected at 28 stations, were identified using electron microscopy (EM). Eight species (P. americana, P. fraudulenta, P. subpacifica, P. heimii, P. pungens, P. seriata, P. delicatissima and P. turgidula), and a novel form, P. sp. GOM, were identified. Species identity was confirmed by sequencing the large subunit of the ribosomal rDNA (28S) and the internal transcribed spacer 2 (ITS2) for six species (36 isolates). Phylogenetic analyses (including neighbor joining, maximum parsimony, and maximum likelihood estimates and ITS2 secondary structure analysis) and morphometric data supported the placement of P. sp. GOM in a novel clade that includes morphologically and genetically similar isolates from Australia and Spain and is genetically most similar to P. pseudodelicatissima and P. cuspidata. Seven species (46 isolates) were grown in nutrient-replete batch culture and aliquots consisting of cells and growth medium were screened by Biosense ASP ELISA to measure total domoic acid (DA) produced (intracellular+extracellular); P. americana and P. heimii were excluded from all toxin analyses as they did not persist in culture long enough for testing. All 46 isolates screened produced DA in culture and total DA varied among species (e.g., 0.04–320 ng ml−1 for P. pungens and P. sp. GOM isolates) and among isolates of the same species (e.g., 0.24–320 ng ml−1 for P. sp. GOM). The 15 most toxic isolates corresponded to P. seriata, P. sp. GOM and P. pungens, and fg DA cell−1 was determined for whole cultures (cells and medium) using ELISA and liquid chromatography (LC) with fluorescence detection (FLD); for seven isolates, toxin levels were also estimated using LC with mass spectrometry and ultraviolet absorbance detection. P. seriata was the most toxic species (up to 3500 fg cell−1) and was observed in the GOM region during all cruises (i.e., during the months of April, May, June and October). P. sp. GOM, observed only during September and October 2007, was less toxic (19–380 fg cell−1) than P. seriata but more toxic than P. pungens var. pungens (0.4 fg cell−1). Quantitation of DA indicated that concentrations measured by LC and ELISA were positively and significantly correlated; the lower detection limit of the ELISA permitted quantification of toxicity in isolates that were found to be non-toxic with LC methods. The confirmation of at least seven toxic species and the broad spatial and temporal distribution of toxic Pseudo-nitzschia spp. have significant implications for the regional management of nearshore and offshore shellfisheries.

Margalef's mandala and phytoplankton bloom strategies☆

Abstract: Margalef's mandala maps phytoplankton species into a phase space defined by turbulence (A) and nutrient concentrations (Ni); these are the hard axes. The permutations of high and low A and high and low Ni divide the space into four domains. Soft axes indicate some ecological dynamics. A main sequence shows the normal course of phytoplankton succession; the r–K axis of MacArthur and Wilson runs parallel to it. An alternative successional sequence leads to the low A-high Ni domain into which many red tide species are mapped. Astronomical and biological time are implicit. A mathematical transformation of the mandala (rotation) links it to the classical bloom models of Sverdrup (time) and Kierstead and Slobodkin (space). Both rarity and the propensity to form red tides are considered to be species characters, meaning that maximum population abundance can be a target of natural selection. Equally, both the unpredictable appearance of bloom species and their short-lived appearances may be species characters. There may be a correlation too between these features and long-lived dormant stages in the life-cycle; then the vegetative planktonic phase is the ‘weak link’ in the life-cycle. Red tides are thus due to species which have evolved suites of traits which result in specific demographic strategies.

New cold-water coral occurrences in the Eastern Ionian Sea: Results from experimental long line fishing

Abstract: Our knowledge on the cold-water corals (CWCs) occurring in the deep waters of Eastern Ionian Sea (E. Mediterranean) has improved with new species records and information related to their occurrence, abundance, distribution, morphological and other characteristics. Data were collected from accidental coral catches during experimental long-line fishing using two different hook sizes (No. 7 targeting hake and No. 9 targeting blackspot seabream). Eight coral taxa were identified, with new records for the E. Ionian Sea for Antipathes dichotoma, Desmophyllum dianthus, Swiftia pallida and Pennatula phosphorea . CWCs occurred in 72% of the long lines and most of them were entire and live (except Leiopathes glaberrima ). The most frequently caught coral group was Antipatharia. Isidella elongata and A. dichotoma presented the highest abundance. The main bulk of the corals was caught between 500 and 600 m depth. Some of the identified corals ( L. glaberrima and I. elongata ) seemed to have more close relationships with some species of the fish assemblage ( Galeus melastomus, Helicolenus dactylopterus ) inhabiting the study area. Species richness and coral catch in numbers were higher for the smaller hook size (No. 9), which caught the larger entire living colonies of A. dichotoma, L. glaberrima and I. elongata . As a result, the corals collected during the present study, characterized by slow growth rates and high longevities, could be considered vulnerable to the gear used in blackspot seabream fishery; this implies the need for more research, monitoring, protection and sustainable management.

Comparison of different wind products and buoy wind data with seasonality and interannual climate variability in the southern Bay of Biscay (2000–2009)

Abstract: Ocean surface winds are essential factors in determining oceanographic and atmospheric processes that can affect ocean circulation and wave generation. Accurate surface wind datasets are needed, therefore, to enable the proper analysis of these processes. Wind data from six databases (National Centers for Environmental Prediction reanalysis (NCEP Reanalysis II), European Centre for Medium-Range Weather Forecasts (ECMWF) re-analysis (ERA-Interim), Modern-Era Retrospective-analysis for Research and Applications (MERRA), NCEP Climate Forecast System Reanalysis (CFSR), QuikSCAT and Cross-Calibrated Multi-Platform (CCMP)) were compared with wind measured in situ by four ocean buoys at the southern limit of the Bay of Biscay. The study covered the period 2000–2009 in such a way that the extent of the time series reduced the margin of error and allowed the disaggregation of the wind data using velocity bins and direction sectors. Statistical results confirmed that datasets with finer spatial resolution (lower than 0.5°×0.5°) gave better results, especially in near-shore areas. A more complete analysis was, therefore, carried out using the finer resolution datasets (QuikSCAT, CCMP and CFSR). This comparison showed that all the datasets were less accurate at low wind speeds ( −1 ) and more accurate at moderate wind speeds. The calculated mean wind speed errors were similar for the three datasets, and the lowest value (1.67 m s −1 ) was from the CCMP dataset. The lowest mean error for wind direction (~37°) was also observed in the CCMP data. The lowest mean wind speed (and direction) bias was obtained from the QuikSCAT data, and the next lowest from the CFSR data. The seasonality for north and east wind components was also determined for the last decade and the results were consistent with forcing for the continental slope current seasonality and winter temperatures or Navidad by wind stress. Correlations between NAO and north and east wind components were low showing that NAO could not be used as a proxy for local wind stress in the southern Bay of Biscay.

Delineating ecological regions in marine systems: Integrating physical structure and community composition to inform spatial management in the eastern Bering Sea

Abstract: Characterizing spatial structure and delineating meaningful spatial boundaries have useful applications to understanding regional dynamics in marine systems, and are integral to ecosystem approaches to fisheries management. Physical structure and drivers combine with biological responses and interactions to organize marine systems in unique ways at multiple scales. We apply multivariate statistical methods to define spatially coherent ecological units or ecoregions in the eastern Bering Sea. We also illustrate a practical approach to integrate data on species distribution, habitat structure and physical forcing mechanisms to distinguish areas with distinct biogeography as one means to define management units in large marine ecosystems. We use random forests to quantify the relative importance of habitat and environmental variables to the distribution of individual species, and to quantify shifts in multispecies assemblages or community composition along environmental gradients. Threshold shifts in community composition are used to identify regions with distinct physical and biological attributes, and to evaluate the relative importance of predictor variables to determining regional boundaries. Depth, bottom temperature and frontal boundaries were dominant factors delineating distinct biological communities in this system, with a latitudinal divide at approximately 60°N. Our results indicate that distinct climatic periods will shift habitat gradients and that dynamic physical variables such as temperature and stratification are important to understanding temporal stability of ecoregion boundaries. We note distinct distribution patterns among functional guilds and also evidence for resource partitioning among individual species within each guild. By integrating physical and biological data to determine spatial patterns in community composition, we partition ecosystems along ecologically significant gradients. This may provide a basis for defining spatial management units or serve as a baseline index for analyses of structural shifts in the physical environment, species abundance and distribution, and community dynamics over time.

Distribution, abundance, biomass and diversity of benthic infauna in the Northeast Chukchi Sea, Alaska: Relation to environmental variables and marine mammals

Abstract: In summer 2009 and 2010, as part of Chukchi Sea Offshore Monitoring in Drilling Area – Chemical and Benthos (COMIDA CAB) program, we performed a quantitative assessment of the biomass, abundance, and community structure of benthic infaunal populations of the Northeastern Chukchi Sea. This analysis documented a benthic species inventory of 361 taxa collected from 142 individual van Veen grab samples (0.1 m−2) at 52 stations. Infaunal abundance was dominated by Polychaeta, Mollusca, and Crustacea. Large concentrations of bivalves (up to 1235 m−2; 920.2 gww m−2) were collected south of Hanna Shoal where flow from two water masses converge and deposit labile carbon to the seafloor, as indicated by low surface sediment C:N ratios. Amphipods (up to 1640 m−2; 26.0 gww m−2), and polychaetes (up to 4665 m−2; 114.7 gww m−2) were documented from multiple stations west of and within Barrow Canyon. This high productivity was most likely due to the “canyon effect”, where marine and coastal detrital carbon supplies are channeled by the canyon structure, enhancing carbon deposition and flux, which supports rich benthic communities within the canyon and surrounding areas. To examine the relationships between infaunal distributions of all collected taxa with the physical environment, we used a Biota and Environment matching (BIO–ENV) routine. A combination of water depth, bottom-water temperature and salinity, surface sediment total organic nitrogen (TON) and sediment C:N molar ratios correlated closest with infaunal abundance distribution (ρ=0.54), indicating that multiple factors influence the success of benthic communities. BIO–ENV routines produced similar correlation results when performed on targeted walrus prey items (bivalves (ρ=0.50), polychaetes (ρ=0.53), but gray whale prey items (amphipods) were not strongly correlated to any combination of physical environmental factors (ρ=0.24). Distributions of primary prey items for gray whales (amphipods) and walruses (bivalves, gastropods and polychaetes) were compared with gray whale and walrus distribution as described by sightings from the 2009 and 2010 aerial survey component of COMIDA. In general, concentrations of walruses and their prey occurred in a swath located south of Hanna Shoal and on the shoal itself although the large differences in sea-ice distribution between the two study years showed that walrus distributions were closely linked to sea-ice location. Other areas within Barrow Canyon and the shelf west of the canyon showed high concentrations of benthic amphipods that were coincident with gray whale sightings as quantified by COMIDA aerial surveys. Overall, data collected on this project indicate that the Northeast Chukchi Sea supports a highly productive and diverse benthic ecosystem that is of significant importance to higher trophic level megafauna.

Indications for algae-degrading benthic microbial communities in deep-sea sediments along the Antarctic Polar Front

Abstract: Phytoplankton blooms in surface waters of the oceans are known to influence the food web and impact microbial as well as zooplankton communities. Numerous studies have investigated the fate of phytoplankton-derived organic matter in surface waters and shelf sediments, however, little is known about the effect of sinking algal biomass on microbial communities in deep-sea sediments. Here, we analyzed sediments of four regions in the Southern Atlantic Ocean along the Antarctic Polar Front that had different exposures to phytoplankton bloom derived organic matter. We investigated the microbial communities in these sediments using high-throughput sequencing of 16S rRNA molecules to determine microorganisms that were active and catalyzed reporter deposition fluorescence in situ hybridization to infer their abundance and distribution. The sediments along the Antarctic Polar Front harbored microbial communities that were highly diverse and contained microbial clades that seem to preferably occur in regions of high primary productivity. We showed that organisms affiliated with the gammaproteobacterial clade NOR5/OM60, which is known from surface waters and coastal sediments, thrive in the deep-sea. Benthic deep-sea NOR5 were abundant, diverse, distinct from pelagic NOR5 and likely specialized on the degradation of phytoplankton-derived organic matter, occupying a similar niche as their pelagic relatives. Algal detritus seemed to not only fuel the benthic microbial communities of large areas in the deep-sea, but also to influence communities locally, as we found a peak in Flavobacteriaceae-related clades that also include degraders of algal biomass. The results strongly suggest that phytoplankton-derived organic matter was rapidly exported to the deep-sea, nourished distinct benthic microbial communities and seemed to be the main energy source for microbial life in the seafloor of vast abyssal regions along the Antarctic Polar Front.

Habitat characterization of deep-water coral reefs in La Gaviera canyon (Avilés Canyon System, Cantabrian Sea)

Abstract: Surveys conducted at the complex Aviles Canyon System (southern Bay of Biscay) in order to identify vulnerable habitats and biological communities revealed the presence of noteworthy deep-water coral reefs in one of the tributaries of the system (La Gaviera Canyon). The aim of the present study is to determine why this deep-sea canyon provides suitable environmental conditions for corals to grow. This hanging canyon is characterized by an irregular U-shaped floor with two narrow differentiated flanks. Sand ripples and rocky outcrops structured in diverse W–E directed steps are observed on the canyon floor, suggesting intense hydrodynamic activity. Accordingly, high-frequency near-bottom current and thermal structure profiles showed that there occur strong shifts in currents/hydrography behaving as front-like features at each tidal cycle. These involve the sudden increase of along-axis velocities to over 50 cm/s and vertical velocities of over 5 cm/s in each tidal cycle associated with the passage of sharp thermal fronts and thermal inversions suggesting overturning. A year-long near-bottom current record showed events with near-bottom velocities well over 1 m/s lasting for several days. Three cold-water coral settings were distinguished: a dense coral reef located on stepped rocky bottoms of the eastern and western flanks, carbonate mounds (20–30 m high) located on the canyon floor, and a cluster of shallower water dead coral framework at the head sector of the canyon. Video and still images from a towed sled and ROV verified the presence of dropstones and rippled sand sheets surrounding the mounds and revealed changes in the coral population (alive or dead; total or patchy coverage) in coral reef and carbonate mound areas. The dominant species of the reef are Lophelia pertusa and Madrepora oculata, which considerably increase the habitat′s complexity and biodiversity in relation to other facies described in the canyon. The presence of living cold-water reefs is directly related to a high-energy environment at depths between 700 and 1200 m in the levels between the lower bound of Eastern North Atlantic Central Water (ENACW) and the core of Mediterranean Water (MW). Such level matches the water density range σθ=27.35–27.65 kg m−3 which has been identified as limits for cold-water coral distribution in the North Atlantic.

Phytoplankton assemblage structure in and around a massive under-ice bloom in the Chukchi Sea

Abstract: Standard and imaging flow cytometry were used to examine the composition of phytoplankton assemblages in and around a massive under-ice bloom in the Chukchi Sea in 2011. In the core of this bloom, roughly 100 km northwest of Hanna Shoal, diatoms represented roughly 87% of the water column carbon-specific biomass of phytoplankton, while nanophytoplankton contributed ~9%. Picoeukaryotes were also observed in this bloom, as were phycoerythrin-containing cells consistent with Synechococcus spp., but picophytoplankton, dinoflagellates, and prymnesiophytes each represented only ~1% of the bloom׳s phytoplankton biomass. More broadly along this part of the Chukchi shelf, nanophytoplankton typically comprised a larger fraction of phytoplankton biomass in the water column, 22% on average but up to 82% at certain locations. Dinoflagellates and prymnesiophytes contributed at most 2% of water column biomass at any location and were most abundant in the deeper slope stations northeast of Hanna Shoal, east of the bloom. Picophytoplankton were most abundant in these deeper slope stations as well, and also in recently ice-free areas to the south around Hanna Shoal. These cell-derived estimates of phytoplankton carbon biomass, which were computed from imaging and standard cytometric observations of phytoplankton cell sizes and from published carbon:volume relationships, agree well with independent measurements of particulate organic carbon concentration from traditional biochemical assays.

Boundary influences on HAB phytoplankton ecology in a stratification-enhanced upwelling shadow

Abstract: Coastal marine ecosystems are profoundly influenced by processes that originate from their boundaries. These include fluid boundaries—with the atmosphere, oceanic boundary currents and terrestrial aquatic systems, as well as solid boundaries—with the seafloor and coast. Phytoplankton populations transfer complexly interacting boundary influences into the biosphere. In this contribution, we apply data from an ocean observing and modeling system to examine boundary influences driving phytoplankton ecology in Monterey Bay, CA, USA. The study was focused on species that may cause harmful algal blooms (HABs). During September–October 2010, autonomous molecular analytical devices were moored at two locations characterized by different degrees of stratification and exposure to upwelling dynamics. The time-series revealed multiple transitions in local HAB phytoplankton communities, involving diatoms (Pseudo-nitzschia spp.), dinoflagellates (Alexandrium catenella), and raphidophytes (Heterosigma akashiwo). Observational and model results showed that the biological transitions were closely related to environmental changes that resulted from a variety of boundary processes—responses of oceanic circulation to wind forcing, influxes of different water types that originated outside the bay, and emergence of strongly stratified nearshore water into the greater bay. Boundary processes were further implicated at patch scales. High-resolution mapping and sampling of a phytoplankton-enriched patch were conducted in a Lagrangian framework using autonomous underwater vehicles. These highly resolved measurements showed that small-scale spatial patterns in the toxicity of Pseudo-nitzschia populations were related to the coupling of resuspended sediments from the bottom boundary layer to the surface mixed layer.

Role of shelfbreak upwelling in the formation of a massive under-ice bloom in the Chukchi Sea

Abstract: In the summer of 2011, an oceanographic survey carried out by the Impacts of Climate on EcoSystems and Chemistry of the Arctic Pacific Environment (ICESCAPE) program revealed the presence of a massive phytoplankton bloom under the ice near the shelfbreak in the central Chukchi Sea. For most of the month preceding the measurements there were relatively strong easterly winds, providing upwelling favorable conditions along the shelfbreak. Analysis of similar hydrographic data from summer 2002, in which there were no persistent easterly winds, found no evidence of upwelling near the shelfbreak. A two-dimensional ocean circulation model is used to show that sufficiently strong winds can result not only in upwelling of high nutrient water from offshore onto the shelf, but it can also transport the water out of the bottom boundary layer into the surface Ekman layer at the shelf edge. The extent of upwelling is determined by the degree of overlap between the surface Ekman layer and the bottom boundary layer on the outer shelf. Once in the Ekman layer, this high nutrient water is further transported to the surface through mechanical mixing driven by the surface stress. Two model tracers, a nutrient tracer and a chlorophyll tracer, reveal distributions very similar to that observed in the data. These results suggest that the biomass maximum near the shelfbreak during the massive bloom in summer 2011 resulted from an enhanced supply of nutrients upwelled from the halocline seaward of the shelf. The decade long trend in summertime surface winds suggests that easterly winds in this region are increasing in strength and that such bloom events will become more common.

Trophodynamics and organic matter assimilation pathways in the northeast Chukchi Sea, Alaska

Abstract: We analyzed trophic linkages in the northeast Chukchi Sea shelf based on the stable carbon and nitrogen isotopic analysis of 39 species collected in 2009 and 2010. To decipher organic matter assimilation pathways, benthic fauna were first categorized into nine trophic guilds based on their physical location in the seabed (epibenthic, surface, or subsurface), feeding mode (suspension feeder, deposit feeder, predator, or scavenger), and food source (suspended particulate organic matter, phytoplankton, zooplankton, sediment, microflora, meiofauna, or macrofauna). A discriminant function analysis (DFA) determined that feeding modes were predicted by stable isotope values at an overall classification success rate of 42%, although classification success of each individual guild varied from 0 to 66%. In some instances, stable isotopes classified trophic guilds incorrectly more often than correctly, suggesting high trophic redundancy in the system. A striking pattern was observed where the δ13C values of individuals in some trophic guilds, ranging from about −23 to −17‰, were substantially more 13C-enriched than representative end-members, which includes phytoplankton (−24.0‰), suspended particulate organic matter (−24.3‰), and bulk sediment organic matter (−23.3‰). In contrast, near-seafloor suspended particulate organic matter was significantly 13C-enriched compared to suspended particulate organic matter of near surface waters (p

Insights into the population dynamics of the deep-sea coral genus Paramuricea in the Gulf of Mexico

Abstract: Species in the genus Paramuricea are among the most common corals in deep waters (>200 m) of the Gulf of Mexico. Paramuricea spp. increase habitat heterogeneity and provide substrate for numerous faunal associates, including ophiuroids that occur on the majority of coral colonies. In light of the Deepwater Horizon oil spill in the Gulf of Mexico, it has become apparent that there is a critical need for data on population dynamics within this genus. To date, at least one species of Paramuricea (P. biscaya) is known to be negatively impacted by the spill. Using remotely operated vehicles from 2009 to 2011, we documented the density and size frequency distributions of Paramuricea across 21 sites at depths of 250–2500 m. Molecular barcoding (mtCOI+igr+MutS) was used to delineate species boundaries among the specimens collected. Results suggested that seven haplotypes are present in the Gulf, and appear to be partially segregated by depth [(type H:

Influence of mesoscale eddies on biological production in the Mozambique Channel: Several contrasted examples from a coupled ocean-biogeochemistry model

Abstract: The impact of mesoscale activity on phytoplankton and nutrient distribution in the Mozambique Channel was simulated by coupling a biogeochemical model (PISCES) with a regional oceanic model (ROMS). Examples of the effects of eddies on the biogeochemistry of the Mozambique Channel are presented to illustrate the complexity of the system. In the model, several cyclonic eddies were found with low concentrations of chlorophyll at their cores, which contrasts with previous studies in the open ocean. In addition, several anticyclonic eddies were simulated with high concentrations of chlorophyll at their cores. Phytoplankton growth within these mesoscale features (both cyclonic and anticyclonic eddies) occurred in response to nutrient injection into the euphotic zone by advection, and subsequent retention of surrounding nutrient-rich waters within eddies. Offshore nutrient distributions depended strongly on lateral advection of nutrient-rich water from the coastal regions, induced by eddy interaction with the shelf. The environmental conditions at the locations where eddies were generated had an important effect on nutrient concentrations within these structures.