210Po and 210Pb in zooplankton fecal pellets

²¹⁰Po and ²¹⁰Pb concentrations in fecal pellets from the zooplanktonic euphausiid Meganyctiphanes norvegica are reported. The ²¹⁰Po:²¹⁰Pb activity ratio is 2.2±0.3, a value in good agreement with that found in suspended particulate matter in surface seawater. Estimates of ²¹⁰Po and ²¹⁰Pb removal times from the mixed layer by fecal pellets alone yield values which are of the same order of magnitude as the removal times for these nuclides by all routes. It is suggested that there is a high probability that zooplanktonic fecal pellets play a significant role in the removal of both these nuclides from the surface layers of the ocean.     

Sinking rates of fecal pellets from gelatinous zooplankton (Salps,Pteropods, Doliolids)

Sinking rates were determined for fecal pellets produced by gelatinous zooplankton (salps, Salpa fusiformis and Pegea socia; pteropods, Corolla spectabilis; and doliolids, Dolioletta gegenbaurii) feeding in surface waters of the California Current. Pellets from the salps and pteropods sank at rates up to 2 700 and 1 800 m d^-1, respectively; such speeds exceed any yet recorded for zooplankton fecal pellets. Fecal pellets of salps were rich in organic material, with C:N ratios from 5.4 to 6.2, close to values for living plankton. The relation between volume and sinking rate indicates that salp and pteropod pellets are slightly less dense than those of pelagic Crustacea; moreover, pellet density varied between different collection dates, probably because of differences in composition. In contrast, doliolid pellets sank at rates up to 208 m d^-1, a rate much lower than would be expected from pellet size. Thus, density and sinking rates of pellets are much more variable in zooplankton than would be expected from studies of crustaceans alone. Moreover, the extraordinarily high sinking rates of fecal pellets of salps indicates that these tunicates may be disproportionately important in the flux of biogenic materials during periods when they form dense population blooms.     

Zooplankton feeding ecology: contents of fecal pellets of the copepods Temora turbinata and T. stylifera from continental shelf and slope waters near the mouth of the Mississippi River

In-situ feeding habits of the copepods Temora turbinata and T. stylifera were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Pellet contents were compared with assemblages of available phytoplankton in the water column at the times of zooplankton sampling. Samples were collected in continental shelf and slope waters of the Gulf of Mexico near the mouth of the Mississippi River. Both species ingested a wide size range and taxonomic array of phytoplankters, and to a lesser extent, other crustaceans. Fecal pellets contained primarily the remains of the phytoplankters that were most abundant in the water at times of collection. There was considerable overlap in the food items ingested by adult females of both copepod species, or two stages of T. turbinata copepodites. Thus, T. turbinata and T. stylifera are omnivores, but primarily opportunistic herbivores.     

Sinking rates of natural copepod fecal pellets

Many pure samples of natural fecal pellets have been collected from mixed small copepods and from the pontellid copepod Anomalocera patersoni in the Ligurian Sea, using a specially designed pellet collection device. Sinking rates of fresh pellets and pellets aged up to 33 days have been determined at 14°C, the mean temperature of the essentially isothermal water column in the Ligurian Sea. Sinking rates of pellets collected during calm sea states increased with increasing pellet volume, but sinking rates of pellets collected during rough sea (Beaufort scale 6) showed little correlation with pellet size. Much of the variability in the sinking rate-pellet size relationships was the result of different pellet composition and compaction, but not pellet age. Pellets produced from laboratory diets of phytoplankton and phytoplankton-sediment mixes showed the expected wide variability in sinking rates, with sediment-ballasted pellets sinking much faster than pellets produced from pure algal diets; thus determination of vertical material fluxes in the sea using laboratory-derived fecal pellet sinking rates is unwarranted. Natural pellet sinking data for small copepods and A. patersoni have been combined with similar data for euphausiids, to yield sinking rates of roughly two orders of magnitude over three orders of magnitude in pellet volume. Pellets from small copepods sank at speeds too slow to be of much consequence to rapid material flux to the deep sea, but they undoubtedly help determine upper water distribution of materials. Recalculation of fecal pellet mass flux estimates from the literature, using our sinking rate data for natural small copepod pellets, yielded estimates about half those of previously published values. Earlier studies had concluded that small fecal pellets were of lesser significance to total material flux than fecal matter; our recalculation strengthens that conclusion. Pellets from large copepods and euphausiids, however, have the capability to transport materials to great depths, and probably do not substantially recycle materials near the surface. The fact that the majority of pellets which had previously been collected in deep traps by other workers were of a size comparable to pellets from our large copepods supports the contention that these larger pellets are the main ones involved in vertical flux.     

A simulation analysis of continental shelf food webs

Energy flow through continental shelf food webs was examined using a simulation model. The model structure expands the two traditional marine food chains of phytoplankton-zooplankton-pelagic fish and benthos-demersal fish into a complex web which includes detritus, dissolved organic matter (DOM), bacteria, protozoa, and mucus net feeders. Simulation of energy flux for different shelf systems using the expanded web revealed that heterotrophic microorganisms and their predators account for a significant component of the energy flux in the continental shelf ecosystem. Contrary to previous models, where all phytoplankton were considered to be grazed by zooplankton, our simulation results indicate that only slightly more than 50% of the annual net primary production is grazed. A substantial quantity of the phytoplankton production directly becomes detritus. Bacteria mineralize detritus and DOM produced by phytoplankton and other components of the food web, converting these to biomass with high efficiency. Consequently, the model predicts that planktonic bacterial production is equivalent to zooplankton production. Exclusion of the bacteria requires the assumption that all DOM is either exported from the system or consumed by another component of the food web. Neither of these assumptions can be supported by present knowledge of the dynamics of DOM in the sea. Model simulations were also employed to test the hypothesis that production exceeds consumption on continental shelves, resulting in exports of 50% of the annual primary production. Simulations of shelves with high rates of primary production resulted in a particulate export of 27% and realistic estimates of secondary production. Results of other simulations suggest that shelves with lower primary production cannot export production and still maintain the macrobenthos and their predators. General properties about continental shelves can also be inferred from the model. From simulations of shelves of differing primary production, nanoplankton are predicted to account for a greater proportion of the primary production in nutrient limited systems. Benthic production appears to be related to both the quantity of primary production and the sinking rates of the phytoplankton. The model indicates that zooplankton fecal inputs to the shelf benthos are only a small portion of the total detrital flux, leading to the prediction that fecal pellets are of little significance in determining benthic production. Finally, the model generates production efficiencies that are highly variable depending on the type of system and kind of populations involved. We argue that the assumed ecological efficiency of 10% should be abandoned for continental shelves and other ecosystems.     

Diet of anguilloid larvae: leptocephali feed selectively on larvacean houses and fecal pellets

Gut contents of 234 leptocephali comprising eight species of eels were examined from five families (Congridae, Muraenidae, Muraenesocidae, Nettastomatidae, and Ophichthidae). The larvae belonged to developing leptocephalus (215 specimens) and the early metamorphic stage (19 specimens). Visible gut contents were recognized in 111 individuals among the eight species, regardless of developmental stage. Two kinds of visible objects, transparent (0.4 to 1.2 mm) and opaque (20 to 380 m), were found in the gut of leptocephali. From their morphological characteristics, the former and the latter were identified as larvacean houses and zooplankton fecal pellets, respectively. Furthermore, most fecal pellets in the gut were identified as oikopleurid larvacean's fecal pellets. No trace of the many other phytoplankton or zooplankton, which were found with leptocephali in the ambient waters and could be suitable-size food, was found in the gut of any leptocephalus. On the basis of the importance of larvecean houses in the diet of several species of leptocephalus larvae, it is proposed that the peculiar, large, fang-like teeth of leptocephali are used for feeding, and evolved to pierce and grasp the mucous houses of larvaceans.     

Production and fate of copepod fecal pellets across the Southern Indian Ocean

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 μg C m⁻³ day⁻¹. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg m⁻² day⁻¹).     

Production,composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as g C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d^-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.     

A time-dependent model of nutrient distribution in continental shelf waters

A time dependent, vertical plane mathematical model of nitrate distribution in Onslow Bay, North Carolina, is developed using similarity theory and conventional numerical techniques. Inputs of nitrate into Onslow Bay are the result of Gulf Stream intrusions (Atkinson et al., 1980) and this forcing is included as a boundary condition for this system. Advective and diffusive processes provide the mechanisms for transport of nitrate in Onslow Bay. The time dependence of the resulting nitrate fields is determined by the rate of phytoplankton removal of nitrate.Nondimensional numbers, arising from model formulation, indicate the relative importance of various processes included in the model. Two nondimensional numbers, H and P, and the nondimensional ratio, P/H, indicate interactions of physical processes. Importance of the biological terms is determined by a third nondimensional number, A. Model results are compared to nitrate data collected in Onslow Bay, North Carolina during an intrusion.     

Increased spatial variance accompanies reorganization of two continental shelf ecosystems

Phase transitions between alternate stable states in marine ecosystems lead to disruptive changes in ecosystem services, especially fisheries productivity. We used trawl survey data spanning phase transitions in the North Pacific (Gulf of Alaska) and the North Atlantic (Scotian Shelf) to test for increases in ecosystem variability that might provide early warning of such transitions. In both time series, elevated spatial variability in a measure of community composition (ratio of cod [Gadus sp.] abundance to prey abundance) accompanied transitions between ecosystem states, and variability was negatively correlated with distance from the ecosystem transition point. In the Gulf of Alaska, where the phase transition was apparently the result of a sudden perturbation (climate regime shift), variance increased one year before the transition in mean state occurred. On the Scotian Shelf, where ecosystem reorganization was the result of persistent overfishing, a significant increase in variance occurred three years before the transition in mean state was detected. However, we could not reject the alternate explanation that increased variance may also have simply been inherent to the final stable state in that ecosystem. Increased variance has been previously observed around transition points in models, but rarely in real ecosystems, and our results demonstrate the possible management value in tracking the variance of key parameters in exploited ecosystems.     

Effects of a decrease in downwelling irradiance on the daytime vertical distribution patterns of zooplankton and micronekton

The daytime vertical distribution of several species of crustaceans, gelatinous zooplankton, and fish were monitored in situ simultaneously with measurements of downwelling irradiance in Oceanographer Canyon in July 1999. During this submersible-based research cruise, an influx of turbid water significantly decreased downwelling irradiance and had a substantial impact on the depth distributions of a number of organisms. Several species of crustaceans, (Thysanoessa gregaria and Sergestes arcticus) and gelatinous zooplankton (Salpa aspera and Salpa fusiformis) ascended over 100 m in the water column during the influx and returned to their pre-influx depths once the influx had ceased. In situ light measurements demonstrated that each of these species was associated with the same irradiance levels during the influx as they were under pre- and post-influx conditions. By contrast, a statistically significant change in temperature, salinity, and oxygen concentrations measured post-influx had no apparent impact on the depth distributions. These results indicate that these species were adjusting their depth distributions to remain within a range of preferred irradiances. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s002270020788     

Sinking rates of fecal pellets from the marine copepod Pontella meadii

Sinking rates of fecal pellets produced by the marine copepod Pontella meadii, grazing on 4 different phytoplankton diets, ranged from 15 to 153 m/day, with a mean of 66 m/day. Sinking rates, in general, were directly related to fecal pellet volumes, but unrelated to the diets used to produce the fecal pellets. There were two-to-threefold variations in sinking rates between fecal pellets of the same volumes, often produced on the same diets. Twenty repetitions of timed sinking of a single fecal pellet revealed sinking rates varying from 33 to 79 m/day, as well as variations in sinking rates within the course of individual descents. It is suggested that copepod fecal pellets are of such small volumes and densities that their sinking rates are subject to microstructural variations in the most carefully controlled water columns. Scanning electron microscope observations revealed lack of structural damage to some of the diatom frustules in the fecal pellets, suggesting that superfluous feeding may have occurred. Thus, the accelerated sinking rates of copepod fecal pellets may provide a mechanism for nutritional enrichment of the deep-sea ecosystem with organic parcels containing incompletely-assimilated plant material.     

Numerical modelling of diel carbon production and zooplankton grazing on the Scotian shelf based on observational data

Using high resolution vertical distributions of chlorophyll and zooplankton, and field observations of photosynthetic parameters, it is shown that on the Scotian Shelf the peak in the vertical profile of primary production generally lies shallower than the chlorophyll maximum, but coincides with the peak in the vertical profile of copepods. A simple numerical model shows that the 24th carbon budget can be balanced using the best available estimates of the rate constants for phytoplankton growth, zooplankton grazing and vertical migration. This calculation is very sensitive to the size of the weight-specific ration and favors values of ～40% d^?1 for it.     

Relation between Engraulis anchoita concentrations and presence of Merluccius hubbsi on the patagonian continental shelf

During the second cruise of the R.V. Professor Siedlecki in spring and summer, 1973, concentrations of anchovy and hake were reported in the area off Golfo Nuevo and the Bay of San Mathias, Argentina. Observations were mainly made with the help of acoustic equipment. The echo recordings revealed an interrelationship between the presence of anchovy schools and the appearance of hake concentrations. This paper demonstrates that this relation is trophic in character.     

Geolocation of free-ranging fish on the European continental shelf as determined from environmental variables

Demersal fish cannot readily be tracked using satellite-based or light-based geolocation techniques. As an alternative, we describe the tidal location method, which uses tidal data recorded by electronic data storage tags (DSTs), to determine geoposition. Times of high water (H) and tidal ranges (R) recorded by DSTs moored at known locations, and from free-swimming tagged plaice, Pleuronectes platessa, were compared with a North Sea tidal database to identify all positions with matching values of H and R. Within the recording precision of the tag (ǂ.2 m, ᆞ min) and the predicted accuracy of the model generated tidal data (ǂ.15 m, ᆨ min), geolocations over much of the North Sea and eastern English Channel were predicted to be accurate to within 40 km, sometimes to within 10 km. Positional estimates of the moored tags were within 15.7Dž.5 km of the actual locations. Geolocations made from tagged plaice within 5 days of release and 5 days pre-recapture were within 35ᆬ km and 37ᆫ km of release and recapture positions respectively. Our results demonstrate the ability of this method to accurately describe the migrations of North Sea plaice throughout their geographical range with a level of accuracy unattainable using light-based geolocation. The method could equally be applied to any shelf-dwelling demersal fish that periodically rests on the sea-bed for the duration of a tidal cycle. In fisheries management, the method has clear potential application in defining the movements and migrations of other commercial species.     

Effects of zooplankton diel vertical migration on a phytoplankton community: A scenario analysis of the underlying mechanisms

A mechanistic model was applied to study the influence of diurnal vertical migration (DVM) of planktonic crustaceans on the succession and composition of the phytoplankton community. While zooplankton was restricted to only one functional group, the phytoplankton community was divided into two functional groups which are distinguished by their maximum growth rates and vulnerability to zooplankton grazing. DVM causes a pulsed grazing regime and may also entail a corresponding reduction of the cumulative daily rates of ingestion and losses of zooplankton. To study the relative importance of these two mechanisms of DVM to phytoplankton we performed a scenario analysis consisting of 5 different scenarios. The results show that DVM has a strong influence on the phytoplankton community. Well edible algae benefit during the first 3–4 weeks of summer stratification by reduced daily grazing. The typical shift from small, well edible algae to larger, poorly or non-edible phytoplankton is distinctly delayed. Under the assumption of unchanged daily grazing, however, a pulsed grazing regime has nearly no influence on the resulting phytoplankton composition. As similar effects are also found for completely non-edible phytoplankton, indirect effects via phosphorus availability must be assumed. Thus, the scenario analysis reveals that the observed effects of DVM on phytoplankton can be explained by a combination of two mechanisms: (1) reduction of the daily zooplankton grazing, and (2) changed assimilation and remineralisation of phosphorus. Surprisingly and in contradiction to earlier reports there is almost no DVM effect on phytoplankton due to the sole action of a pulsed grazing regime.     

Megabenthic decapod crustacean assemblages on the Galician continental shelf and upper slope (north-west Spain)

The structure of megabenthic decapod crustacean assemblages on the Galician (north-west Spain) continental shelf (100 to 200 depth) and upper slope (200 to 500 m) was analyzed based on surveys carried out in autumn and spring, from 1980 to 1987. Forty species belonging to 19 families were caught. The portunid crab Polybius henslowii, a species with pelagic phases, was the most abundant species, but displayed strong spatial and temporal fluctuations. Other dominant species were the Norway lobster Nephrops norvegicus, the portunid Liocarcinus depurator, the galatheids Munida intermedia and M. sarsi, and the shrimps Solenocera membranacea, Plesionika heterocarpus, Pasiphaea sivado and Dichelopandalus bonnieri. Total abundance and biomass (average values excluding Polybius henslowii = 255 individuals and 2.06 kg/30 min tow) and species richness and diversity, H′ (6.85 species and H′ = 1.45 per tow) displayed a significant positive correlation with depth, and strong interannual fluctuations. The factors determining community organization were depth and, to a lesser extent, spatial structure. There was clear evidence of bathymetric zonation, differentiating between species characteristic of the slope (D. bonnieri and Pasiphaea sivado), shelf-slope edge (Macropipus tuberculatus, Pontophilus spinosus, Munida sarsi, S. membranacea, Processa spp.) and shelf (L. depurator, Macropodia tenuirostris, Paguridae and Chlorotocus crassicornis). The spatial zonation was related to changes in oceanography and sediment along the continental margin. Goneplax rhomboides, N. norvegicus, C. crassicornis and Alpheus glaber are benthic species which generally exhibit burrowing behaviour, and they were found mainly in the southern area where there are fine sediments due to the outwelling from the Rías Baixas. Different benthopelagic shrimps (Pontophilus spinosus, Plesionika heterocarpus,Processa spp. and Pasiphaea sivado) were typical of the zone just north of Fisterra, characterized by a convergence of water masses bringing about an increase in productivity due to upwelling. The benthic anomuran and brachyuran crabs Munida intermedia, M.␣sarsi, L.␣depurator and Macropipus tuberculatus were characteristic of the northwestern zone between Fisterra and Estaca, where the infauna reaches high biomass despite coarser sediments with a lower concentration of organic material than in the southern area. Lastly, both the Paguridae and Macropodia tenuirostris were species typically found in the waters in the northern shelf. Based on interannual changes in assemblage structure, two periods could be distinguished: between 1980 and 1984, when Polybius henslowii, D. bonnieri and Pasiphaea sivado had abundance peaks; and another period from 1985 to 1987 when L. depurator, Munida intermedia, M.␣sarsi and Macropipus tuberculatus increased in abundance. Received: 21 May 1996 / Accepted: 9 August 1996     

Sea ice retreat alters the biogeography of the Bering Sea continental shelf.

Seasonal ice cover creates a pool of cold bottom water on the eastern Bering Sea continental shelf each winter. The southern edge of this cold pool, which defines the ecotone between arctic and subarctic communities, has retreated approximately 230 km northward since the early 1980s. Bottom trawl surveys of fish and invertebrates in the southeastern Bering Sea (1982-2006) show a coincident reorganization in community composition by latitude. Survey catches show community-wide northward distribution shifts, and the area formerly covered by the cold pool has seen increases in total biomass, species richness, and average trophic level as subarctic fauna have colonized newly favorable habitats. Warming climate has immediate management implications, as 57% of variability in commercial snow crab (Chionoecetes opilio) catch is explained by winter sea ice extent. Several measures of community distribution and structure show linear relationships with bottom temperature, suggesting warming climate as the primary cause of changing biogeography. However, residual variability in distribution not explained by climate shows a strong temporal trend, suggesting that internal community dynamics also contribute to changing biogeography. Variability among taxa in their response to temperature was not explained by commercial status or life history traits, suggesting that species-specific responses to future warming will be difficult to predict.     

Domoic acid in benthic flatfish on the continental shelf of Monterey Bay,California, USA

Within Monterey Bay, California, USA, the food web transfer of domoic acid (DA), a neurotoxin produced by diatoms of the genus Pseudo-nitzschia, has led to major mortality events of marine mammals and birds. Less visible, and less well known, is whether invertebrates and fish associated with the benthos are also affected by blooms of DA-producing Pseudo-nitzschia spp. This study examines the presence of DA in benthic flatfish offshore of Davenport, California, (37°0′36″N, 122°13′12″W) and within Monterey Bay, California (36°45′0″N, 122°1′48″W), including species that feed primarily in the sediment (benthic-feeding) and species that feed primarily in the water column (benthopelagic-feeding). Flatfish caught between 10 December 2002 and 17 November 2003 at depths of 30–180 m had concentrations of DA in the viscera ranging from 3 to 26 μg DA g⁻¹ of viscera. Although the DA values reported are relatively low, benthic-feeding flatfish were frequently contaminated with DA, especially as compared with the frequency of contamination of flatfish species that feed in the water column. Furthermore, on days in which both benthic-feeding and benthopelagic-feeding flatfish were collected, the former had significantly higher concentrations of DA in the viscera. Curlfin turbot, Pleuronicthys decurrens, the flatfish with both the highest level and frequency of DA contamination, are reported to feed exclusively on polychaetes, suggesting that these invertebrates may be an important vector of the toxin in benthic communities and may pose a risk to other benthic-feeding organisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.