Life history of the deep-sea cephalopod family Histioteuthidae in the western Mediterranean

The life cycle of the two species of the deep-sea family Histioteuthidae inhabiting the Mediterranean Sea (Histioteuthis reversa and Histioteuthis bonnellii) was studied from monthly samples taken throughout the year during daytime hours by bottom trawl gears. A small sample of individuals found floating dead on the sea surface was also analyzed. Both species were caught exclusively on the upper slope at depths greater than 300 m. Their frequency of occurrence increased with depth and showed two different peaks, at 500–600 m and 600–700 m depth in H. bonnellii and H. reversa, respectively, which might indicate spatial segregation. Maturity stages were assigned using macroscopic determination and confirmed with histological analyses. Although mature males were caught all year round, no mature females were found, which suggests that their sexual maturation in the western Mediterranean takes place deeper than the maximum depth sampled (800 m). In fact, the increase in mean squid size with increasing depth in H. reversa indicates an ontogenetic migration to deeper waters. The individuals of both species found floating dead on the sea surface were spent females which had a relatively large cluster of small atresic eggs and a small number of remaining mature eggs scattered in the ovary and mantle cavity. The sizes of these females were clearly larger than the largest individuals caught with bottom trawls. A total of 12 and 7 different types of prey, belonging to three major taxonomic groups (crustaceans, osteichthyes and cephalopods), were identified in the stomach contents of H. reversa and H. bonnellii, respectively. In both species fishes were by far the main prey followed by crustaceans, whereas cephalopods were found only occasionally. The preys identified, mainly myctophids and natantian crustaceans, indicate that both histioteuthids base their diet on pelagic nictemeral migrators.     

Life history of the bathyal octopus Pteroctopus tetracirrhus (Mollusca,Cephalopoda) in the Mediterranean Sea

The life cycle of the deep-sea octopus Pteroctopus tetracirrhus was studied from monthly samples obtained throughout the year in different areas of the western Mediterranean (mainly around the Balearic Islands and along the coast of the Iberian Peninsula). A total of 373 individuals (205 females, 168 males) were analyzed; females ranged from 4.5 to 14.0 cm mantle length (ML) and males from 4.5 to 11.5 cm ML. There were few small-sized octopuses (<7 cm ML) in the samples, which might indicate that these individuals inhabit rocky grounds that are not accessible to trawlers or waters deeper than the maximum depth sampled (800 m). The species occurred more frequently around the Balearic Islands than along the Iberian Peninsula as they appeared in 20% and 7%, respectively, of the hauls in these areas. The octopus inhabits the lower continental shelf and upper slope in both areas, primarily between 200 and 500 m depth. Modal lengths were followed from autumn, when recruits were caught by trawlers, to summer, when reproduction took place. Females grew from 8 to 10 cm ML from winter to spring, but this modal size did not increase further in summer; males grew from 7 to 9 cm ML from winter to spring. The total disappearance of large individuals after summer suggests a life cycle lasting a single year. The evolution of the monthly mean sizes showed that the growth was best described by log-linear functions in both sexes. The length at first maturity was clearly higher in females (12 cm ML) than in males (8 cm ML). A total of 30 different prey items, belonging to four major taxonomic groups (crustaceans, osteichthyes, cephalopods and gastropods), were identified in the stomach contents. The diet of the octopus was based on crustaceans and teleosts, which accounted for 75% and 23% of the prey items, respectively. Cephalopods and gastropods were accessory prey as they only represented 1.6% and 0.7%, respectively, of the total. The octopus showed a marked preference for the benthic fish Symphurus nigrescens and the endobenthic crustacean Alpheus glaber. The bathymetric distribution of P. tetracirrhus coincides with those of these two main prey, which suggests that the distribution of the octopus might be strongly linked to its trophic resources.     

Influence of summer conditions on the larval fish assemblage in the eastern coast of Tunisia (Ionian Sea,Southern Mediterranean)

The structure of the summer larval fish assemblage off the eastern coast of Tunisia and its relation to environmental conditions was studied, from ichthyoplankton samples taken during a survey conducted between 23rd June and 9th July 2008. A total of 68 larval fish taxa were identified, 52 to species level. The taxonomic composition and abundance of the larval fish assemblage showed high spatial heterogeneity. Mesoscale hydrographic features, such as eddies, seem to play an important role in the spatial distribution of fish larvae in the area, enhancing concentration and retention. The larval fish assemblage was dominated by the small pelagic species Sardinella aurita (26.6% of the total larval fish abundance), followed by Engraulis encrasicolus (22.6%), Spicara spp. (8.6%) and Mullus barbatus (6.8%). Shannon–Weaver index (H′) ranged between 0 and 2.62. The highest values were found offshore, at 95 miles east of Sousse, over depths around 250 m. The diversity was higher in this region as a result of transport by currents and retention by eddies. It has also been shown that the eastern coast of Tunisia is a spawning ground for the tuna species Auxis rochei, Thunnus thynnus and Thunnus alalunga. Larvae of mesopelagic fishes represented 5.46% of the total abundance, with Cyclothone braueri, Ceratoscopelus maderensis and Lampanyctus crocodilus being the most important species. Canonical correspondence analysis (CCA) indicated that depth was the most important environmental factor in explaining species distribution.     

The feeding habits of slope dwelling macrourid fishes in the eastern North Pacific

The diet of slope dwelling macrourid fishes in the eastern North Pacific is poorly known. We collected several hundred stomach samples to investigate the feeding habits of Coryphaenoides acrolepis and Albatrossia pectoralis, the two dominant slope dwelling macrourids off the continental United States. Coryphaenoides acrolepis exhibited a pronounced ontogenetic shift in diet. Specimens <15 cm pre-anal fin length (PAF) consumed primarily polychaetes, amphipods, cumaceans and mysids, while larger individuals consumed increasingly larger, more pelagic prey such as fish, squid, and large crustaceans. Scavenging was also very important to specimens >15 cm with scavenged food constituting approximately 20% of the weight of total prey and occurring in approximately 20% of fish 21–29 cm. Albatrossia pectoralis consumed primarily midwater fish and squid, and we believe that it feeds in the water column. There were significant differences between the diets of A. pectoralis and C. acrolepis suggesting some degree of niche separation between macrourid species on the continental slope of the eastern North Pacific. Both species are at the top of the food web on the upper continental slope and, because of their abundance, may exert significant pressures on their prey populations.     

Distribution and diversity of open-ocean,near-bottom macroplankton in the western Mediterranean: Analysis at different spatio-temporal scales

We analyzed the distribution, diversity, and composition of western Mediterranean macroplankton (excluding gelatinous taxa) in the water column over depths of ca. 550–850 m, with special attention to near-bottom (0–1.5 and ca. 5–77 m above the bottom, mab) levels, and including data from three areas (off the coasts of Catalonia, and to the NW, and SE of Mallorca, Balearic Islands) in the period 1991–2008. Spatio-temporal changes in macroplankton abundance were evaluated as follows: (i) by seasonal sampling in 2007 off the Catalonian coast, (ii) by comparing Catalonian and Balearic Island slopes, and (iii) by comparing a fixed station on the Catalonian slope (at 550–800 m depth) at decadal (1991/1992–2007/2008) time scales. Diversity (in terms of species richness, S) was greater (i) at ca. 5–77 mab than at 0–1.5 mab, (ii) over the insular slopes of the Balearic Island (around Mallorca) than over the mainland Catalonian slopes, and (iii) in the period 1991/1992 than in 2007, likely related to higher values of the North Atlantic Oscillation (NAO) index in 1991/1992. In most analyses species composition was strongly influenced by the degree of stratification and homogenization of the water column in summer–autumn and winter–spring respectively and by location (longitude). Changes consisted mainly of higher density of macroplankton (e.g. abundance of the dominant euphausiids Nematoscelis megalops, Meganyctiphanes norvegica and Euphausia krohni and of the fish Cyclothone braueri) between June and October, parallel to an increase in the T and S close to the bottom. This coincided with changes in the flow of Levantine intermediate water (LIW) in the area. Aggregation of adult forms of the dominant species close to the bottom in summer–autumn could be favored because summer is the period of highest density of food – copepods, mainly Calanus helgolandicus – near the bottom off the Catalan slope. The formation of a thermocline in the water column and the reinforcement of the permanent thermohaline front at the shelf-slope break during summer at ca. 400 m in the Balearic Basin may also enhance this tendency toward greater aggregation of deep macroplankton under stratified water column conditions.     

Characterization of the solar light field within the ocean mesopelagic zone based on radiative transfer simulations

The solar light field within the ocean from the sea surface to the bottom of the mesopelagic zone was simulated with a radiative transfer model that accounts for the presence of inelastic radiative processes associated with Raman scattering by water molecules, fluorescence of colored dissolved organic matter (CDOM), and fluorescence of chlorophyll-a contained in phytoplankton. The simulation results provide a comprehensive characterization of the ambient light field and apparent optical properties (AOPs) across the entire visible spectral range within the depth range 200–1000 m of the entire mesopelagic zone for varying chlorophyll-a concentration and seawater optical properties in the mixed surface layer of the ocean. With increasing depth in the mesopelagic zone, the solar irradiance is reduced by ~9–10 orders of magnitude and exhibits a major spectral maximum in the blue, typically centered around a light wavelength of 475 nm. In the green and red spectral regions, the light levels are significantly lower but still important owing to local generation of photons via inelastic processes, mostly Raman scattering and to a lesser extent CDOM fluorescence. The Raman scattering produces a distinct secondary maximum in irradiance spectra centered around 565 nm. Comparisons of our results with light produced by the radioactive decay of the unstable potassium isotope contained in sea salt (⁴⁰K) indicates that the solar irradiance dominates over the ⁴⁰K-produced irradiance within the majority of the mesopelagic zone for most scenarios considered in our simulations. The angular distribution of radiance indicates the dominance of downward propagation of light in the blue and approach to uniform distribution in the red throughout the mesopelagic zone. Below the approximate depth range 400–500 m, the shape of the angular distribution is nearly invariant with increasing depth in the green and red and varies weakly in the blue. The AOPs at any light wavelength also assume nearly constant values within the deeper portion of the mesopelagic zone. These results indicate that the mesopelagic light field reaches a nearly-asymptotic regime at depths exceeding ~400–500 m.     

Factors influencing the abundance of deep pelagic bioluminescent zooplankton in the Mediterranean Sea

Measurements of the density of deep pelagic bioluminescent zooplankton (BL) were made with the Intensified Silicon Intensifier Target (ISIT) profiler in the Ligurian, Tyrrhenian, Adriatic, Ionian Seas and the Strait of Sicily from ～300 m to near seafloor. Mean BL densities ranged from 2.61 m^?3 at 500–1000 m depth in the Adriatic Sea to 0.01 m^?3 at 4000–5000 m depth in the E Ionian Sea. We investigated drivers of spatial variation of deep pelagic BL density. Linear regression was applied between surface chlorophyll a (Chl a) concentration and underlying BL density. Chl a values were determined from satellite derived 100 km radius composites (six 10-day means per ISIT deployment, over preceding 60 days). At 500–1000 m depth we found a significant positive relationship between mean BL density and mean Chl a in the period prior to 0–10 days (at 1% level) and prior to 10–40 days (at 5% level). Beyond 40 days no relationship between BL density and Chl a was found at this depth. At depths 1000–1500 m BL density values were low and no significant relationship with Chl a was detected. Generalised additive modelling (GAM) was used to assess the influence of benthic hotspots (seamount; cold water coral mound; mud volcano) on overlying BL density. A reduction in BL density was found downstream of the Palinuro seamount from 300 to 600 m. No effect on BL density in the overlying water column was detected from the presence of cold water corals. Higher BL densities were detected over the W Madonna dello Ionio mud volcano than at other sites sampled in the NW Ionian Sea. We find surface Chl a to be a good predictor of BL density in the mesopelagic zone; below this depth we hypothesise that processes affecting the efficiency of particle export to deep water may exert greater influence on BL density.     

Greenland halibut diet in the Northwest Atlantic from 1978 to 2003 as an indicator of ecosystem change

The Northwest Atlantic marine community underwent dramatic changes during the last 30 years, including the collapse of many groundfish stocks and an increase in shrimp populations. Greenland halibut Reinhardtius hippoglossoides is an important commercial species and one of the top fish predators in this system. It is a large, wide-ranging flatfish that is found at depths up to 2200 m and it has an opportunistic diet which makes it a potential candidate for an ecosystem indicator. Analysis of stomach contents of Greenland halibut between 1978 and 2003 indicates that diet composition reflects the major changes in community structure. Over the entire period there was a clear increase in the importance of invertebrates, particularly after 1992. This change was associated with a higher importance of Pandalus shrimp and Gonatus squid and a protracted reliance on zooplankton by predators under 25 cm length. Capelin Mallotus villosus was the dominant prey between 1978 and 1992 for predators in the 12–63 cm range, but its importance dropped off drastically in the mid 1990s. Levels of main prey in the diet of Greenland halibut correlated well with fishery-independent surveys. Greenland halibut sample capelin well, compared to bottom trawl surveys and acoustic surveys. Greenland halibut consumed small shrimp which are not routinely caught by surveys and may be important in deriving information on year classes and growth of shrimp. Our results suggest that Greenland halibut's diet is a useful tracker of ecosystem change.     

Bioenergetic model estimates of interannual and spatial patterns in consumption demand and growth potential of juvenile pink salmon (Oncorhynchus gorbuscha) in the Gulf of Alaska

A bioenergetic model of juvenile pink salmon (Oncorhynchus gorbuscha) was used to estimate daily prey consumption and growth potential of four ocean habitats in the Gulf of Alaska during 2001 and 2002. Growth potential was not significantly higher in 2002 than in 2001 at an alpha level of 0.05 (P=0.073). Average differences in growth potential across habitats were minimal (slope habitat=0.844 g d⁻¹, shelf habitat=0.806 g d⁻¹, offshore habitat=0.820 g d⁻¹, and nearshore habitat=0.703 g d⁻¹) and not significantly different (P=0.630). Consumption demand differed significantly between hatchery and wild stocks (P=0.035) when examined within year due to the interaction between hatchery verses wild origin and year. However, the overall effect of origin across years was not significant (P=0.705) due to similar total amounts of prey consumed by all juvenile pink salmon in both study years. We anticipated that years in which ocean survival was high would have had high growth potential, but this relationship did not prove to be true. Therefore, modeled growth potential may not be useful as a tool for forecasting survival of Prince William Sound hatchery pink salmon stocks. Significant differences in consumption demand and a two-fold difference in nearshore abundance during 2001 of hatchery and wild pink salmon confirmed the existence of strong and variable interannual competition and the importance of the nearshore region as being a potential competitive bottleneck.     

Clearance rates of ephyrae and small medusae of the common jellyfish Aurelia aurita offered different types of prey

Prey selection and knowledge of the amounts of water processed by the early stages of the common jellyfish Aurelia aurita may at certain times of the year be crucial for understanding the plankton dynamics in marine ecosystems with mass occurrences of this jellyfish. In the present study we used two different methods (“clearance method” and “ingestion-rate method”) to estimate the amount of water cleared per unit of time of different types and sizes of prey organisms offered to A. aurita ephyrae and small medusae. The mean clearance rates of medusae, estimated with Artemia sp. nauplii as prey by both methods, agreed well, namely 3.8 ± 1.4 l h^? 1 by the clearance method and 3.2 ± 1.1 l h^? 1 by the ingestion-rate method. Both methods showed that copepods (nauplii and adults) and mussel veligers are captured with considerably lower efficiency, 22 to 37% and 14 to 30%, respectively, than Artemia salina nauplii. By contrast, the water processing rates of ephyrae measured by the clearance method with A. salina nauplii as prey were 3 to 5 times lower than those measured by the ingestion-rate method. This indicates that the prerequisite of full mixing for using the clearance method may not have been fulfilled in the ephyrae experiments. The study demonstrates that the predation impact of the young stages of A. aurita is strongly dependent on its developmental stage (ephyra versus medusa), and the types and sizes of prey organisms. The estimated prey-digestion time of 1.3 h in a steady-state feeding experiment with constant prey concentration supports the reliability of the ingestion-rate method, which eliminates the negative “container effects” of the clearance method, and it seems to be useful in future jellyfish studies, especially on small species/younger stages in which both type and number of prey can be easily and precisely assessed.     

The habitat of mesopelagic scyphomedusae in Monterey Bay,California

In the mesopelagic zone, at depths of 200–1000 m in the Monterey Submarine Canyon, CA, medusae in three genera of scyphozoa, Atolla, Periphylla and Poralia, were observed, videotaped and collected over a 9-year period (1990–1998). Environmental data were obtained simultaneously using a remotely operated vehicle (ROV) with sensors for depth, temperature, salinity and dissolved oxygen. Shipboard measurements of these same properties at two reference stations in the region defined the local water masses and helped identify species niches using the metric of spiciness and oxygen levels of the waters in which medusae were visually “captured”. The most abundant genus of mesopelagic scyphomedusae was Atolla, found associated most strongly with the spicy (warm, salty) waters of the California Undercurrent, usually above the core of the oxygen minimum zone (OMZ; O₂>0.5 ml/l). The least abundant mesopelagic scyphomedusa was Periphylla, which occurred in more variable waters, including those with a greater contribution of fresher, colder (less spicy), subarctic water and, hence, most like those at the offshore California Current station in the most depleted oxygen zone (averaging O₂ <0.3 ml/l). Poralia was mostly confined to the densest, coldest water, with peak abundance at the lower boundary of the OMZ (i.e., 0.3< O₂<0.5 ml/l). These spiciness measures on local isopycnal surfaces within the mesopelagic zone, supported by data on dissolved oxygen concentrations, indicate highly significant but fine-scale habitat differences in species habitats in Central California waters. This in situ investigation appears to be one of only a few studies to document fine-scale, water mass affinities of mesopelagic zooplankton.     

Trophic relationships at intrannual spatial and temporal scales of macro and megafauna around a submarine canyon off the Catalonian coast (western Mediterranean)

The spatial and temporal changes of near-bottom macrofauna (suprabenthos and macroplankton) and the trophic relationships of megabenthic decapod crustaceans were analyzed off the Catalonian coasts (western Mediterranean) around Berenguera submarine canyon in four periods (April and December 1991, March and July 1992) and four zones (within Berenguera Canyon at ca. 450 m, and on adjacent slope at ca. 400, 600 m and 1200 m). In March 1992, we found the highest macrofauna abundance and the smallest sizes in the canyon, suggesting a positive effect of river discharges on suprabenthos recruitment. By contrast, macroplankton (decapods, fishes and euphausiids) did not show higher recruitment into canyons. After analyzing the diet of 23 decapod crustaceans, we found a significant segregation between guilds feeding on zooplankton and on benthos. Zooplankton (euphausiids and Pasiphaeidae) and infauna (polychaetes, Calocaris macandreae and ophiuoroids) were consistently the main prey exploited by decapod crustaceans around Berenguera Canyon. We also found some macrophyte (Posidonia oceanica) consumption, which was higher in periods of water column homogeneity (winter–spring and late autumn). Positive correlations between decapods' gut fullness (F) and decapod abundance indicate feeding aggregations, while positive correlations were also found between F and Llobregat River (situated ca. 18 km from Berenguera head) flow 1 to 2 months before sampling. Increases in F were delayed only 1 month when zooplankton feeders were analyzed alone, while benthos feeders did not show significant relationships with any environmental variables. That indicates that the response of megabenthic decapods feeding on benthos to environmental shifts is slower than that of zooplankton feeders. The importance of river flows in enhancing food supply of macro- and megabenthos dwelling close to submarine canyons was apparent, with a delay in the fauna response of 0–2 months after river flow peaks.     

Paleoenvironment changes in the NW Okhotsk Sea for the last 18 kyr determined with micropaleontological,geochemical, and lithological data

Lithological, geochemical, and micropaleontological data indicate that the Late Glacial of the northwestern Okhotsk Sea (OS) is characterised by severe climatic and environmental conditions with mainly perennial sea ice coverage and low productivity accompanied by weak deep-water ventilation and a temperate formation of the upper Sea of Okhotsk Intermediate Water (SOIW). The age model of the studied core sediments was constructed by AMS ¹⁴C dating. The most severe environmental conditions occurred during the period 15.8–14.8 kyr, synchronous with cold Heinrich event 1. Insignificant regional environmental amelioration accompanied by an increase of productivity and ice weakening during summer occurred almost simultaneously with the Bølling–Allerøed (BA) warming. The obtained results distinguished both the Bølling and Allerøed warmings as having different environmental conditions. Oxygen content in the surface sediment was low, as seen from the production of the benthic foraminifera (BF) species. During 12.6–11.1 kyr, synchronous with the Younger Dryas (YD) cold event, the regional environment conditions were cold, but not as severe as the glacial ones. Some climatic warming since the Preboreal has stimulated sea ice melting and surface amelioration during the summer season, which in turn led to a productivity rise and changes in the water column and bottom environment. Some increase in the surface water stratification and the intensified oceanic diatom and surface radiolarian production is parallel with the development of a mesopelagic regime of productivity. The surface sediment condition favours BF abundance and domination by BF species tolerant to oxygen deficiencies. During the Boreal period more stable surface conditions were accompanied by continuously high productivity and an intensifying of its mesopelagic regime.Significant regional climate warming since the Atlantic (9 kyr ago) strongly intensified the summer sea ice melting in the OS, and this created considerable surface environment amelioration with the preferential transport of bacteria and phytodetritus into the SOIW. Further considerable warming of the regional climate from 6 kyr ago contributed to slight sea ice changes, surface water warming, and the enhancement of its stratification; all typical for most of the OS. Along with a high nutrient supply from the Amur River, the NW OS experienced a strong diatom production increase with the maximum amount occurring during the last 3.6 kyr. This changed the productivity type and organic matter export into the water column while increasing the feeding of the “productive” Plagoniidae spp. group and decreasing the microbial biomass supply into the upper SOIW. Some sea surface water cooling or saltier conditions at the beginning of the Subatlantic (2.4–1.8 kyr) was followed by its warming or freshening 1.5–1.0 kyr ago, which likely correlated with the Medieval Warm Period. In turn, that probably led to strong surface water stratification, productivity deterioration and considerable changes in the overall NW OS environment. The established sequence of the northwestern OS environmental changes during the Late Glacial–Holocene is related to the Northern Hemispheric climate changes and was likely forced by atmospheric teleconnection in line with the polar circulation index variability.     

Feeding ecology of mesopelagic zooplankton of the subtropical and subarctic North Pacific Ocean determined with fatty acid biomarkers

Mesopelagic zooplankton may meet their nutritional and metabolic requirements in a number of ways including consumption of sinking particles, carnivory, and vertical migration. How these feeding modes change with depth or location, however, is poorly known. We analyzed fatty acid (FA) profiles to characterize zooplankton diet and large particle (>51 μm) composition in the mesopelagic zone (base of euphotic zone ?1000 m) at two contrasting time-series sites in the subarctic (station K2) and subtropical (station ALOHA) Pacific Ocean. Total FA concentration was 15.5 times higher in zooplankton tissue at K2, largely due to FA storage by seasonal vertical migrators such as Neocalanus and Eucalanus. FA biomarkers specific to herbivory implied a higher plant-derived food source at mesotrophic K2 than at oligotrophic ALOHA. Zooplankton FA biomarkers specific to dinoflagellates and diatoms indicated that diatoms, and to a lesser extent, dinoflagellates were important food sources at K2. At ALOHA, dinoflagellate FAs were more prominent. Bacteria-specific FA biomarkers in zooplankton tissue were used as an indicator of particle feeding, and peaks were recorded at depths where known particle feeders were present at ALOHA (e.g., ostracods at 100–300 m). In contrast, depth profiles of bacterial FA were relatively constant with depth at K2. Diatom, dinoflagellate, and bacterial biomarkers were found in similar proportions in both zooplankton and particles with depth at both locations, providing additional evidence that mesopelagic zooplankton consume sinking particles. Carnivory indices were higher and increased significantly with depth at ALOHA, and exhibited distinct peaks at K2, representing an increase in dependence on other zooplankton for food in deep waters. Our results indicate that feeding ecology changes with depth as well as by location. These changes in zooplankton feeding ecology from the surface through the mesopelagic zone, and between contrasting environments, have important consequences for the quality and quantity of organic material available to deeper pelagic and benthic food webs, and for organic matter sequestration.     

Environmental drivers of megafaunal assemblage composition and biomass distribution over mainland and insular slopes of the Balearic Basin (Western Mediterranean)

The influence of mesoscale physical and trophic variables on deep-sea megafauna, a scale of variation often neglected in deep-sea studies, is crucial for understanding their role in the ecosystem. Drivers of megafaunal assemblage composition and biomass distribution have been investigated in two contrasting areas of the Balearic basin in the NW Mediterranean: on the mainland slope (Catalonian coasts) and on the insular slope (North of Mallorca, Balearic Islands). An experimental bottom trawl survey was carried out during summer 2010, at stations in both sub-areas located between 450 and 2200 m water depth. Environmental data were collected simultaneously: near-bottom physical parameters, and the elemental and isotopic composition of sediments. Initially, data were analysed along the whole depth gradient, and then assemblages from the two areas were compared. Analysis of the trawls showed the existence of one group associated with the upper slope (US=450–690 m), another with the middle slope (MS=1000–1300 m) and a third with the lower slope (LS=1400–2200 m). Also, significant differences in the assemblage composition were found between mainland and insular slopes at MS. Dominance by different species was evident when the two areas were compared by SIMPER analysis. The greatest fish biomass was recorded in both areas at 1000–1300 m, a zone linked to minimum temperature and maximum O₂ concentration on the bottom. Near the mainland, fish assemblages were best explained (43% of total variance, DISTLM analysis) by prey availability (gelatinous zooplankton biomass). On the insular slope, trophic webs seemed less complex and were based on vertical input of surface primary production. Decapods, which reached their highest biomass values on the upper slope, were correlated with salinity and temperature in both the areas. However, while hydrographic conditions (temperature and salinity) seemed to be the most important variables over the insular slope, resource availability (gelatinous zooplankton and Calocaris macandreae) predominated and explained 59% of decapod assemblage variation over the mainland slope. Both fish and decapods were linked to net primary production recorded over the mainland 3 months before sampling, while the delay between the input of food from the surface and fish abundance was only 1 month on the insular slope. Our results suggest that trophic relationships over insular slopes probably involve a shorter food chain than over mainland slopes and one that is likely more efficient in terms of energy transfer.     

Full-depth profiles of prokaryotes,heterotrophic nanoflagellates,and ciliates along a transect from the equatorial to the subarctic central Pacific Ocean

Studies in epipelagic waters report higher heterotrophic microbial biomass in the productive high latitudes than in the oligotrophic low latitudes; however, biogeographical data are scarce in the deep ocean. To examine the hypothesis that the observed latitudinal differences in heterotrophic microbial biomass in the epipelagic zone also occur at depth, abundance and biomass of heterotrophic prokaryotes, nanoflagellates (HNF), and ciliates were determined at depths of 5–5000 m in the central Pacific between August and September of 2005. Heterotrophic microbial biomass increased from the tropical to the subarctic region over the full water column, with latitudinal differences in prokaryotic biomass increasing from 2.3-fold in the epipelagic zone to 4.4-fold in the bathypelagic zone. However, the latitudinal difference in HNF and ciliate biomass decreased with depth. In the mesopelagic zone, the vertical attenuation rate of prokaryotic abundance, which was calculated as the linear regression slope of log-log plot of abundance versus depth, ranged from –0.55 to –1.26 and was more pronounced (steeper slope) in the lower latitudes. In contrast, the vertical attenuation rate of HNF in the mesopelagic zone (–1.06 to –1.27) did not differ with latitude. In the subarctic, the attenuation rate of HNF was 1.7 times steeper than for prokaryotes. These results suggest the accumulation of prokaryotes in the deep subarctic Pacific, possibly due to low grazing pressure. Although the vertical attenuation rate of ciliates was steepest in the bathypelagic zone, HNF abundance did not further decrease at depths below 1000 m, except for at 2000 m where HNF was lowest across the study area. Ciliate abundance ranged 0.3–0.8 cells l^–1 at 4000 m, and were below the detection limit (<0.1 cells l ^–1) at 5000 m. To our knowledge, this study presents the first data for ciliates below 2000 m.     

Bioaccumulation and trophic transfer of mercury in striped bass (Morone saxatilis) and tautog (Tautoga onitis) from the Narragansett Bay (Rhode Island,USA)

We examined the bioaccumulation and trophic transfer of mercury in two marine finfish species, striped bass (Morone saxatilis) and tautog (Tautoga onitis), collected from the Narragansett Bay (Rhode Island, USA). For each of these target fish, white muscle tissue was analyzed for total mercury (Hg) and results were evaluated relative to fish age, body size, and Hg content of preferred prey. Dietary and stable isotope analysis was also used to elucidate the effect of trophic processes on Hg concentrations in fish. The Hg content of muscle tissue was positively correlated with fish age and length for both species, although striped bass accumulated Hg faster than tautog. Accelerated Hg bioaccumulation in striped bass is consistent with its high trophic level (trophic level = 4.07) and Hg-enriched prey (forage fish and macrocrustaceans; mean Hg content = 0.03 mg Hg kg wet wt^?1). In contrast, tautog maintain a lower trophic status (trophic level = 3.51) and consume prey with lower Hg levels (mussels and crabs; mean Hg content = 0.02 mg Hg kg wet wt^?1). Despite differences in Hg bioaccumulation between target fish, the mean Hg concentration of tautog exceeded levels in striped bass (0.24 and 0.16 mg Hg kg wet wt^?1, respectively) due to a disparity in age-at-catch between sampled groups (mean age of tautog and bass = 11.3 and 4.3 yr, respectively). Taking into account legal minimum catch lengths further revealed that 75.0% of legal-size striped bass (>70.2 cm TL; n = 4) and 44.8% of tautog (>40.6 cm TL; n = 29) had Hg levels beyond the US EPA regulatory threshold of 0.3 mg Hg kg wet wt^?1. Moreover, Hg-length relationships suggest that each target fish meets this threshold near their minimum legal catch length. Our findings reiterate the value of species ecology to improve predictions of fish Hg and permit better management of human contamination by this important dietary source.     

Mesozooplankton prey preference and grazing impact in the western Arctic Ocean

The role of mesozooplankton as consumers and transformers of primary and secondary production in the Beaufort and Chukchi Seas was examined during four cruises in spring and summer of both 2002 and 2004 as part of the western Arctic Shelf–Basin Interactions (SBI) program. Forty-seven grazing experiments using dominant mesozooplankton species and life stages were conducted at locations across the shelf, slope, and basin of the Chukchi and Beaufort Seas to measure feeding rates on both chlorophyll and microzooplankton and to determine mesozooplankton prey preferences.Mesozooplankton biomass was at all times dominated by life stages of four copepod taxa: Calanus glacialis, Calanus hyperboreus, Metridia longa, and Pseudocalanus spp. Significant interannual, seasonal, regional, between species and within species differences in grazing rates were observed. Overall, the dominant zooplankton exhibited typical feeding behavior in response to chlorophyll concentration that could be modeled using species and life-stage specific Ivlev functions. Microzooplankton were preferred prey at almost all times, with the strength of the preference positively related to the proportion of microzooplankton prey availability. Average mesozooplankton grazing impacts on both chlorophyll standing stock (0.6±0.5% d⁻¹ in spring, 5.1±6.3% d⁻¹ in summer) and primary production (12.8±11.8% d⁻¹ in spring, 27.6±24.5% d⁻¹ in summer) were quite low and varied between shelf, slope, and basin. Coincident microzooplankton grazing experiments [Sherr, E.B., Sherr, B.F., Hartz, A.J., 2009. Microzooplankton grazing impact in the Western Arctic Ocean. Deep-Sea Research II] were conducted at most stations. Together, microzooplankton–mesozooplankton grazing consumed only 44% of the total water-column primary production, leaving more than half directly available for local export to the benthos or for offshore transport into the adjacent basin.     

Evidence of marine mammal predation of the European eel (Anguilla anguilla L.) on its marine migration

Temperature and depth logging tags were implanted into adult eels released on Atlantic west coasts of France and Ireland to study their oceanic migration behavior. For three of the tags, 25 to 256 days after release there was a dramatic rise in temperature from 10 °C to 36 °C and the dive profile changed from depths of 300–1000 m to repeated ascents to the surface. This indicated that the eels carrying the tags had been eaten by a mammalian predator. Two of the tags had sufficient sampling rate to resolve the dives in detail. They recorded a total of 91 dives to maximum depths of 250–860 m lasting 11–12 min and with surface intervals of 5–7 min. More than two thirds of the dives included a rapid descent from approximately 500 m to 600–700 m. From this we infer that the predator was most likely a deep-diving toothed whale. The dives logged while the tags were inside the predator revealed that the temperature usually decreased during dives, and increased again during surface periods. The temperature drops during dives were probably caused by the ingestion of prey or water. These observations provide insights into the behavior of toothed whales foraging in the mesopelagic zone.     

Parameterization of nonlinear shallow water waves over sloping bottoms

Investigation of the bottom slope effects on the nonlinear transformation of irregular waves, which are generated based on JONSWAP spectra, is carried out in a physical wave flume with three slopes (β = 1/15, 1/30, 1/45). The slope effects on the estimation of representative wave height are examined first. To obtain a better estimation of wave height, the slope effect should be considered when slope is larger than 1/30. The nonlinear parameters (bicoherence, skewness and asymmetry) are estimated by using the wavelet-based bispectrum, and the empirical formulae regarding these nonlinear parameters as a function of the local Ursell number are derived based on the present data measured on each slope. The results indicate that the slopes have a negligible effect on the variations of the skewness. The fitted coefficients of the formulae for the other parameters on slope β = 1/15 are clearly different from the results on the slopes β = 1/30 and 1/45, indicating that slope influence on the parameterization cannot be ignored when β > 1/30. Hence, new formulae considering the slope effect are presented. Furthermore, the empirical formulae for the data in surf zone are recommended.