Spatial spring distribution of the copepod Eurytemora affinis (Copepoda,Calanoida) in a restoring estuary,the Scheldt (Belgium)

The spatial spring distribution of Eurytemora affinis (adults and C5) in the Scheldt estuary (Belgium) brackish and freshwater reaches was studied in between 1996 and 2007. The bulk of the E. affinis population being generally situated in the brackish water reach (salinity > 0.5); we studied which environmental factors are responsible for its recent sporadic occurrence in the freshwater estuarine reach. Using PLS analysis, it is shown that its presence upstream is limited by a sufficient oxygen concentration (>4 mg l⁻¹) that is associated with temperature. Not only are the environmental conditions in the upstream zone important, but also the frequent presence of an O₂ minimum zone in the mid-estuary (O₂ min < 1.3 mg l⁻¹) seems to block the movement of the downstream E. affinis population in an upstream direction. Occasionally, the bulk of the population is however situated upstream. During these periods, high E. affinis abundance was also observed in the Durme tributary. Our findings suggest the possibility to use E. affinis as an “indicator” species of water quality, but also lead us to stress the necessity to consider conditions over the entire estuary when studying restoration effects, not exclusively in the zone of interest.     

Fitting a predator–prey model to zooplankton time-series data in the Gironde estuary (France): Ecological significance of the parameters

The relationships between the seasonal fluctuations of the copepod Eurytemora affinis and the mysid Neomysis integer were studied from observed data and experimental results, using a predator–prey model in the oligo-mesohaline area of the Gironde estuary. Mean seasonal fluctuations of abundances were derived from long term data series collected from 1978 to 2003 for both species. In situ predator–prey experiments over a seasonal cycle were used to estimate the seasonal variation of the consumption rate of N. integer on E. affinis and to verify the order of magnitude of the biological parameters given by the model.Predator–prey experiments revealed a high seasonal variation in maximum consumption rates with a mean of 56 ± 9 ind. pred⁻¹ d⁻¹. Maximum consumption rates were always higher for adults than for juveniles of Neomysis integer. Recorded selectivities were higher on nauplii than on copepodids + adults of Eurytemora affinis, both for the juveniles and the adults of N. integer. Neomysis integer mainly fed on meroplanktonic larvae, when they were available in higher abundances, than E. affinis in their environment.Spring increases of abundance for Eurytemora affinis copepodids + adults seemed to be mainly controlled by temperature whereas its decreasing abundance in summer was more related to Neomysis integer predation, suggesting that summer fluctuations of E. affinis abundance are probably controlled by mysid predation at summer times. Using a Lotka–Volterra predator–prey model, the seasonal peak of abundance of the mysid N. integer was well reproduced considering a predation on copepodids + adults of E. affinis, and suggested a dependence between mysid and copepod seasonal variations. However, the seasonal peak amplitude could not be explained solely by a predation on copepodids + adults or on nauplii of the copepod. Thus, N. integer is probably dependent on the seasonal fluctuations of the copepod's abundance, complementing its diet with macrophytal detritus during periods of scarce food.     

Plankton community diversity from bacteria to copepods in bloom and non-bloom conditions in the Celtic Sea in spring

The plankton community composition comprising heterotrophic bacteria, pro-/eukaryotes, heterotrophic nanoflagellates, microzooplankton and mesozooplankton was assessed during the spring bloom and at non-bloom stations in the English Channel and Celtic Sea between 6 and 12 April 2002. Non-bloom sites were characterised by a dominance of pro-/eukaryotic phytoplankton <20 μm, higher abundance of heterotrophic nanoflagellates, microzooplankton standing stocks ranging between 60 and 380 mg C m⁻², lower mesozooplankton diversity and copepod abundance of between 760 and 2600 ind m⁻³. Within the bloom, the phytoplankton community was typically dominated by larger cells with low abundance of pro-/eukaryotes. Heterotrophic nanoflagellate cell bio-volume decreased leading to a reduction in biomass whereas microzooplankton biomass increased (360–1500 mg C m⁻²) due to an increase in cell bio-volume and copepod abundance ranged between 1400 and 3800 ind m⁻³. Mesozooplankton diversity increased with an increase in productivity. Relationships between the plankton community and environmental data were examined using multivariate statistics and these highlighted significant differences in the abiotic variables, the pro-/eukaryotic phytoplankton communities, heterotrophic nanoflagellate, microzooplankton and total zooplankton communities between the bloom and non-bloom sites. The variables which best described variation in the microzooplankton community were temperature and silicate. The spatial variation in zooplankton diversity was best explained by temperature. This study provides an insight into the changes that occur between trophic levels within the plankton in response to the spring bloom in this area.     

Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

Abundance,community composition,size distribution,and production rates of tintinnids in Narragansett Bay,Rhode Island

The abundance of tintinnid ciliates in lower Narragansett Bay was measured at weekly intervals over the period 1980–1982. Twenty-nine species representing nine genera ranged in abundance from 10¹ to 10⁵ tintinnids l⁻¹. Tintinnopsis was the most numerous genus in terms of numbers of species and individuals. Total abundance increased with water temperature above 6°C, and with nanoplankton chlorophyll a (<10 μm and <5 μm chl a) averaged over the water column. Exceptions occurred during blooms of phytoplankton previously demonstrated to be poor food for tintinnids (Olisthodiscus, Thalassiosira). Tintinnids aggregated near the bottom during periods of low nanoplankton chl a. Abundances and distributions were not correlated with particulate organic carbon. Seasonal changes in length and oral diameter of loricas of individual species were inversely related to temperature. Oral diameter was a more constant feature of lorica morphology than length. Seasonal patterns in the mean oral diameter of the lorica of all species reflected decreases in the oral diameter of individual species with increasing temperature, and changes from small to large species coincident with similar shifts in the size of phytoplankton. The production rate of tintinnids was 3·3 mgCl⁻¹ year⁻¹, equivalent to the ingestion of 26% of total annual net primary production and 52% of the estimated production of <10 μm phytoplankton.     

Environmental control of mesozooplankton community structure in the Seine estuary (English Channel)

This paper is the first to describe the spatio-temporal changes of mesozooplankton in the Seine estuary. Monthly samples were collected along the estuary in 1996 in order to analyse the seasonal changes of the mesozooplankton community and to identify the major environmental parameters that may influence the spatial distribution of zooplankton in this megatidal estuary. Statistical analysis (canonical correspondence analysis) showed that salinity was the main factor correlated with the longitudinal distribution of zooplankton. Marine species (Temora longicornis, barnacle larvae…) were located in the outer part of the estuary, while more oligohaline species (Eurytemora affinis) were recorded in the inner part of the estuary. A mixed zone was characterised by the presence of the neritic copepods Acartia spp. and Eurytemora affinis. The marine species (e.g. T. longicornis, Oikopleura dioica, Barnacle larvae) showed maximum abundance at the end of spring (June) while the most abundant estuarine species, E. affinis, peaked in late winter-spring and declined with the onset of summer. This copepod dominated the estuarine zooplankton throughout the year, and found in the Seine estuary very high favourable conditions to exhibit ultimate abundances (> 190 000 ind m^–3) which is one order of magnitude higher than those found in other European estuaries. It represented the main prey for major planktonivorous species such as suprabenthic and fish species located living in the upstream zone of the Seine estuary.     

Diel vertical migration of the copepod Calanus sinicus before and during formation of the Yellow Sea Cold Bottom Water in the Yellow Sea

To understand the effects of the Yellow Sea Cold Bottom Water(YSCBW)on the diel vertical migration(DVM)of the copepod Calanus sinicus,we surveyed vertical distribution of C.sinicus at a fixed station in the Yellow Sea before(spring)and during(summer)formation of the YSCBW.Cold water(<10 C)was observed in the bottom layer when the water column was thermally stratified in summer,but the water column was thermally well-mixed in spring 2010.Samples were collected from five different layers at 3-h intervals using an opening-closing net.Adult females(1–155 ind./m3)showed a clear normal DVM pattern throughout the entire water column in spring,whereas adult males did not migrate.DVM of copepodite V(CV)individuals was not clear,but the maximum abundance of CI–CIV occurred consistently in the upper 10–20 m layer,where there was a high concentration of chlorophyll-a(Chl-a)(0.49–1.19μg/L).In summer,weak DVM was limited to cold waters beneath the thermocline for adult females(<30 ind./m3),but not for adult males.The maximum abundance of CI–CIV also occurred consistently in the subsurface layer(20–40 m)together with high concentrations of Chl-a(0.81–2.36μg/L).CV individuals(1–272 ind./m3)moved slightly upward nocturnally to the near-surface layer(10–20 m),where the average temperature was 25.74 C,but they were not found in the surface layer(0–10 m;28.31 C).These results indicate that the existence of the YSBCW affected food availability at depth and the vertical temperature distribution,leading to variation in the amplitude and shape of stage-specific vertical distributions(CI to adults)in C.sinicus before and during the formation of cold waters in the Yellow Sea during the study period.     

Grazing by meso- and microzooplankton on phytoplankton in the upper reaches of the Schelde estuary (Belgium/The Netherlands)

In contrast with the marine reaches of estuaries, few studies have dealt with zooplankton grazing on phytoplankton in the upper estuarine reaches, where freshwater zooplankton species tend to dominate the zooplankton community. In spring and early summer 2003, grazing by micro- and mesozooplankton on phytoplankton was investigated at three sites in the upper Schelde estuary. Grazing by mesozooplankton was evaluated by monitoring growth of phytoplankton in 200 μm filtered water in the presence or absence of mesozooplankton. In different experiments, the grazing impact was tested of the calanoïd copepod Eurytemora affinis, the cyclopoid copepods Acanthocyclops robustus and Cyclops vicinus and the cladocera Chydorus sphaericus, Moina affinis and Daphnia magna/pulex. No significant grazing impact of mesozooplankton in any experiment was found despite the fact that mesozooplankton densities used in the experiments (20 or 40 ind. l⁻¹) were higher than densities in the field (0.1–6.9 ind. l⁻¹). Grazing by microzooplankton was evaluated by comparing growth of phytoplankton in 30 and 200 μm filtered water. Microzooplankton in the 30–200 μm size range included mainly rotifers of the genera Brachionus, Trichocerca and Synchaeta, which were present from 191 to 1777 ind. l⁻¹. Microzooplankton had a significant grazing impact in five out of six experiments. They had a community grazing rate of 0.41–1.83 day⁻¹ and grazed up to 84% of initial phytoplankton standing stock per day. Rotifer clearance rates estimated from microzooplankton community grazing rates and rotifer abundances varied from 8.3 to 41.7 μl ind.⁻¹ h⁻¹. CHEMTAX analysis of accessory pigment data revealed a similar phytoplankton community composition after incubation with and without microzooplankton, indicating non-selective feeding by rotifers on phytoplankton.     

Meso- and microzooplankton responses to an in situ iron fertilization experiment (SEEDS II) in the northwest subarctic Pacific

A mesoscale iron fertilization experiment was carried out in the western subarctic Pacific during summer 2004. The iron-patch was traced for 26 days after the enrichment, and the abundance and behavior of meso- and microzooplankton was compared with those outside of the patch. The surface chlorophyll-a concentration in the patch was high between days 10 and 13 (2.5 mg m⁻³) and decreased to the initial level after day 20. Microzooplankton grazing rates, estimated by a dilution method, was mostly balanced with phytoplankton growth rates throughout the observed period. Dominant mesozooplankton species in the upper 200 m were copepods: dominated by Eucalanus bungii, Neocalanus plumchrus and Metridia pacifica. Species composition did not change in the patch over the observation period. The copepod biomass was 3–5 times higher than in Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS), the previous iron-enrichment experiment in the same area, before the bloom, and exponentially increased both inside and outside the patch, which was mainly brought by the development of N. plumchrus. The development rates of N. plumchrus were not significantly different between inside and outside the patch. Estimated grazing rate suggest that the copepod grazing was main cause of the low accumulation of phytoplankton biomass, and dominance of grazing-resistant organisms such as large ciliates, large diatoms and diatoms with extremely long setae. “Arrested migration” for M. pacifica and upward shift of vertical distribution by E. bungii were observed during the bloom period, even if the accumulation of phytoplankton biomass was very low compared to other iron-enrichment experiments. These results indicate that the copepod grazing shaped the food-web structure of the lower trophic levels (biomass and species composition) in SEEDS II.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

Calanoid copepod resting egg abundance and hatching success in the sediment of the Seine estuary (France)

Benthic, viable resting eggs of calanoid copepods were found for the first time in the Seine estuary (France) during July 2008. Vertical distribution of the resting eggs in the sediment was determined up to 10 cm depth. Hatching success of the eggs extracted from different 1-cm thick sediment layers was experimentally tested immediately after extraction and after a long refractory phase (i.e. 11 months) of storage at low temperature (4–5 °C). The hatching success of resting eggs obtained immediately after sediment incubation was lower (0.72%) than the value observed after 11 months (4.50%) with an overall hatching success of 2.37%. The marine, calanoid copepod Temora longicornis was the primary species to hatch from the eggs; however, the estuarine calanoid copepod Eurytemora affinis also hatched from resting eggs. The mean abundance of eggs found in sediment (1.42 × 10⁶ eggs m⁻²) was comparable to that reported for other marine and estuarine calanoid copepods. The Seine estuary sediment had a high variability of egg abundance (between 0.14 and 8.10 × 10⁷ eggs m⁻³) suggesting that the hydrodynamics of this macrotidal estuary are likely responsible for this variability. Significant sediment resuspension occurs in the Seine estuary during flood periods and spring tides leading to resting eggs to contribute along the year to the nauplii recruitment of calanoid copepods. On average, around 400,000 nauplii m⁻³ month⁻¹ of the main calanoid copepods can emerge from the surface layer sediment in the Seine estuary, suggesting that resting eggs could play an important role in the population dynamics of key calanoid copepods in the Seine estuary.     

Seasonal Variations in Abundance and Size Composition of the Lobate Ctenophore Bolinopsis Mikado (Moser) in Tokyo Bay, Central Japan

Surveys of the abundance and size composition of the ctenophore Bolinopsis mikado were conducted in Tokyo Bay over a 5-year period from 1990 to 1994. B. mikado appeared throughout the year, and its mass occurrence was observed between August and November; annual maximum abundance ranged from 19 to 91 ind. m^–2. Water temperature seems to influence the seasonal variation of B. mikado abundance. Environmental conditions (e.g. rough waters due to a typhoon) and predation by the beroid ctenophore Beroe cucumis appear to affect annual variations of B. mikado abundance. Size frequency distributions of B. mikado indicated that its reproduction was most active in summer and fall but occurred throughout the year in Tokyo Bay. A sharp decline of the copepod population in August 1990 was probably due to predation by B. mikado which was very abundant at that time; its predatory impact was estimated to be 24 % day^–1.     

Effects of mesoscale phytoplankton variability on the copepods Neocalanus flemingeri and N. plumchrus in the coastal Gulf of Alaska

The copepods Neocalanus flemingeri and N. plumchrus are major components of the mesozooplankton on the shelf of the Gulf of Alaska, where they feed, grow and develop during April–June, the period encompassing the spring phytoplankton bloom. Satellite imagery indicates high mesoscale variability in phytoplankton concentration during this time. Because copepod ingestion is related to food concentration, we hypothesized that phytoplankton ingestion by N. flemingeri and N. plumchrus would vary in response to mesoscale variability of phytoplankton. We proposed that copepods on the inner shelf, where the phytoplankton bloom is most pronounced, would be larger and have more lipid stores than animals collected from the outer shelf, where phytoplankton concentrations are typically low. Shipboard feeding experiments with both copepods were done in spring of 2001 and 2003 using natural water as food medium. Chlorophyll concentration ranged widely, between 0.32 and 11.44 μg l⁻¹ and ingestion rates varied accordingly, between 6.0 and 627.0 ng chl cop⁻¹ d⁻¹. At chlorophyll concentrations<0.50 μg l⁻¹, ingestion is always low, <40 ng cop⁻¹ d⁻¹. Intermediate ingestion rates were observed at chlorophyll concentrations between 0.5 and 1.5 μg l⁻¹, and maximum rates at chlorophyll concentrations>1.5 μg l⁻¹. Application of these feeding rates to the phytoplankton distribution on the shelf allowed locations and time periods of low, intermediate and high daily feeding to be calculated for 2001 and 2003. A detailed cross-shelf survey of body size and lipid store in these copepods, however, indicated they were indistinguishable regardless of collection site. Although the daily ingestion of phytoplankton by N. flemingeri and N. plumchrus varied widely because of mesoscale variability in phytoplankton, these daily differences did not result in differences in final body size or lipid storage of these copepods. These copepods efficiently dealt with small and mesoscale variations in their food environment such that mesoscale structure in phytoplankton did not affect their final body size.     

Particulate suspended matter concentrations in the Bahía Blanca Estuary,Argentina: Implication for the development of phytoplankton blooms

The inner zone of the Bahía Blanca Estuary is shallow, nutrient-rich and turbid. Tidal energy and water turbulence strongly affect the water column resulting in a well-mixed structure and high concentrations of suspended sediment. The phytoplankton community is mostly dominated by diatoms and the annual pattern has been characterized by a recurrent winter-early spring bloom. Here, we investigated to what extent the temporal variations of suspended particulate matter (SPM) regulate the phytoplankton blooms in the head of the estuary by light-limitation. Sampling was done on a fortnightly basis (weekly during the blooming season) at a fixed station in the inner zone of the estuary from January 2007 to February 2008. SPM concentrations and light extinction coefficients (k) in the water column were significantly correlated and showed relatively lower values during the phytoplankton maximal biomass levels. During winter, SPM and k reached values of 23.6 mg l⁻¹ and 0.17 m⁻¹ which were significantly lower than the annual means of 77.6 mg l⁻¹ and 2.94 m⁻¹, respectively. The particulate organic matter (POM) concentration was significantly correlated with the calculated phytoplankton biomass although the contribution of the latter to the total POM was rather low. Both, POM and biomass, had maximal values during winter (21.8 mg l⁻¹ and 393.5 μg C l⁻¹) and mid summer (24.3 mg l⁻¹ and 407.0 μg C l⁻¹), with cell densities up to 8 × 10⁶ cells l⁻¹ and chlorophyll a up to 24.6 μg l⁻¹. Our results suggest that the decrease of SPM concentrations in the water column with a concomitant increase in the penetration of solar radiation seems to be one of the main causes for the development of the phytoplankton winter bloom in the Bahía Blanca Estuary.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

Vertical distribution of phytoplankton biomass,production and growth in the Atlantic subtropical gyres

Ninety-four stations were sampled in the Atlantic subtropical gyres during 10 cruises carried out between 1995 and 2001, mainly in boreal spring and autumn. Chlorophyll a (Chl-a) and primary production were measured during all cruises, and phytoplankton biomass was estimated in part of them. Picoplankton (<2 μm) represented >60% of total Chl-a concentration measured at the surface, and their contribution to this variable increased with depth. Phytoplankton carbon concentrations were higher in the upper metres of the water column, whereas Chl-a showed a deep maximum (DCM). At each station, the water column was divided into the upper mixed layer (ML) and the DCM layer (DCML). The boundary between the two layers was calculated as the depth where Chl-a concentration was 50% of the maximum Chl-a concentration. On average DCML extends from 67 to 126 m depth. Carbon to Chl-a (C:Chl-a) ratios were used to estimate phytoplankton carbon content from Chl-a in order to obtain a large phytoplankton carbon dataset. Total C:Chl-a ratios averaged (±s.e.) 103±7 (n=22) in the ML and 24±4 (n=12) in the DCML and were higher in larger cells than in picoplankton. Using these ratios and primary production measurements, we derived mean specific growth rates of 0.17±0.01 d⁻¹ (n=173) in the ML and 0.20±0.01 d⁻¹ (n=165) in the DCML although the differences were not significant (t-test, p>0.05). Our results suggest a moderate contribution of the DCML (43%) to both phytoplankton biomass and primary production in the Atlantic subtropical gyres.     

‘ Global Change ’ Impact of Inter-Annual Variation in Water Discharge as a Driving Factor to Dredging and Spoil Disposal in the River Rhine System and of Turbidity in the Wadden Sea

Between 1970 to 2000, the annual mean suspended matter (SPM) concentrations in the Vlie and Marsdiep tidal inlets of the Wadden Sea varied over five times. The present paper examines the possible relationship between SPM in the Wadden Sea and changing river Rhine discharges and dredging operations. The major short-term variations in annual mean SPM in part of the Wadden Sea appears to be a non-linear, exponential, function of river Rhine discharge and dredge spoil disposal (110 km to over 200 km from the area in front of the Dutch coast near the river Rhine outlet). Correlation coefficients (with SPM as the fixed and dredge disposal as the independent variable) ranged from R=0·8 (deep tidal inlet of Marsdiep) to R=0·2 (shallow inner area of Vlie) and weakened mainly as a function of distance to the disposal site. The best correlation with river discharge was for Marsdiep tidal inlet (r=0·45), indicating the superior effect of dredge disposal over river discharge-related processes. Taking the estimated regression equation as an explorative model, indicates that, without any disposal of dredge spoil, the expected SPM concentration levels in the tidal inlets of the Wadden Sea will be <15 g m⁻³ (comparable to the 1950s). The overall mean (and the highest mean) annual concentrations for the investigation period reached 42 (90) g m⁻³ at Marsdiep and 35 (75) g m⁻³ at Vlie. Assuming a 10% (220 m⁻³ s⁻¹) increase in river Rhine discharge over the next 50 years, and unchanged dredging policy and other circumstances, SPM concentrations would increase 5–15% for Marsdiep and Vlie. Compared with the calculated (12·4 g m⁻³ SPM in Marsdiep and 14·8 g m⁻³ SPM in Vlie) and measured (15 g m⁻³ SPM in Marsdiep) background SPM concentrations, the expected overall mean increase since 1950 is at least 250% of background. The natural variation in river Rhine discharge will cause further inter-annual variation. Changes in SPM concentrations, due to expected changes in wind climate, combined with river discharge are estimated to increase SPM concentrations 20% above the present situation. The possible implications of changes in land use in combination with further increasing river discharge, changed wind fields and increased temperature are discussed. An important management conclusion is that increasing mean river discharge will significantly increase the need for dredging and spoil disposal, and result in further elevated SPM concentrations in the Dutch coastal zone and the Wadden Sea. The presented relationships offer possibilities for developing new management strategies in relation to dredging and its effects.     

First distribution record of the invasive copepod Oithona davisae Ferrari and Orsi, 1984, in the coastal waters of the Aegean Sea

The occurrence of the invasive nonindigenous copepod Oithona davisae Ferrari and Orsi, 1984, is reported for the first time in the Aegean Sea. The data we collected in August 2017 from 14 stations along the Turkish coast of the Aegean Sea reveal the spatial distribution of O. davisae between the openning of the Dardanelles Strait in the north and the Izmir Bay in the south. The O. davisae individuals, in seven mesozooplankton samples collected from a single station, were consistently found in the inner part of the Izmir Bay from April 2015‐October 2016. The abundance of female O. davisae ranged from 4 ind./m³ in April 2015 to 31,524 ind./m³ in July 2016 and contributed to the total oithonid female population by 10.8% in April 2015 and 92.8% in September 2016. Our results show that this species is well established in the inner part of Izmir Bay and that it has become a permanent component of the copepod community in the area.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.