Long-term monitoring of the sedimentary processes in the central part of Sagami Bay, Japan: rationale, logistics and overview of results

Deep-sea benthic ecosystems are mainly sustained by sinking organic materials that are produced in the euphotic zone. “Benthic-pelagic coupling” is the key to understanding both material cycles and benthic ecology in deep-sea environments, in particular in topographically flat open oceanic settings. However, it remains unclear whether “benthic-pelagic coupling” exists in eutrophic deep-sea environments at the ocean margins where areas of undulating and steep bottom topography are partly closely surrounded by land. Land-locked deep-sea settings may be characterized by different particle behaviors both in the water column and in relation to submarine topography. Mechanisms of particle accumulation may be different from those found in open ocean sedimentary systems. An interdisciplinary programme, “Project Sagami”, was carried out to understand seasonal carbon cycling in a eutrophic deep-sea environment (Sagami Bay) with steep bottom topography along the western margin of the Pacific, off central Japan. We collected data from ocean color photographs obtained using a sea observation satellite, surface water samples, hydrographic casts with turbidity sensor, sediment trap moorings and multiple core samplings at a permanent station in the central part of Sagami Bay between 1997 and 1998. Bottom nepheloid layers were also observed in video images recorded at a real-time, sea-floor observatory off Hatsushima in Sagami Bay. Distinct spring blooms were observed during mid-February through May in 1997. Mass flux deposited in sediment traps did not show a distinct spring bloom signal because of the influence of resuspended materials. However, dense clouds of suspended particles were observed only in the spring in the benthic nepheloid layer. This phenomenon corresponds well to the increased deposition of phytodetritus after the spring bloom. A phytodetrital layer started to form on the sediment surface about two weeks after the start of the spring bloom. Chlorophyll-a was detected in the top 2 cm of the sediment only when a phytodetritus layer was present. Protozoan and metazoan meiobenthos increased in density after phytodetritus deposition. Thus, “benthic-pelagic coupling” was certainly observed even in a marginal ocean environment with undulated bottom topography. Seasonal changes in features of the sediment-water interface were also documented.     

Particle dynamics in the deep water column of Sagami Bay, Japan. II: seasonal change in profiles of suspended phytodetritus

Vertical distributions of turbidity & phytodetritus (Chl.a and pheopigment), and their seasonal variations were measured in the deep water column of Sagami Bay, Japan, in June 1999, February 2000 and May 2000. Observations were carried out at eight stations along an east-west section of Sagami Bay using a CTD/water sampling system equipped with a memory-type infrared back-scattering meter which had been calibrated for the suspended particles collected in Sagami Bay. Turbidity increased close to the bottom in both summer and winter, indicating the existence of a benthic nepheloid layer throughout the year. But the vertical gradient of turbidity was much larger in summer than in winter. The concentration of Chl.a and pheopigment also increased in the benthic layer in summer, sometimes reaching values of more than 0.01 and 0.2 μg/l, respectively, much higher than those reported in hemipelagic regions of the ocean. In winter, on the other hand, Chl.a kept a constant low value throughout the deep water column. This indicates that the turbid water mass formed in the benthic layer in summer derives from the deposition of large amounts of phytodetritus in spring and the resuspension of these aggregates, which are subsequently decomposed in the benthic layer during the following autumn. Unlike the benthic boundary layer, the turbidity of intermediate water was lower in summer rather than in winter. Because the phytoplankton aggregates exported from the surface water during the spring bloom not only supply phytodetritus to the benthic layer but also scavenge the suspended particles in the water column, the steep vertical gradient of turbidity observed in summer may reflect the dynamic interaction between suspended and sinking particles in the deep water column.     

Material supply to the abyssal seafloor in the Northeast Atlantic

Downward particle flux was measured using sediment traps at various depths over the Porcupine Abyssal Plain (water depth 4850 m) for prolonged periods from 1989 to 1999. A strong seasonal pattern of flux was evident reaching a maximum in mid-summer. The composition of the material changed with depth, reflecting the processes of remineralisation and dissolution as the material sank through the water column. However, there was surprisingly little seasonal variation in its composition to reflect changes in the biology of the euphotic zone.Currents at the site have a strong tidal component with speeds almost always less than 15 cm/sec. In the deeper part of the water column they tend to be northerly in direction, when averaged over periods of several months.A model of upper ocean biogeochemistry forced by meteorology was run for the decade in order to provide an estimate of flux at 3000 m depth. Agreement with measured organic carbon flux is good, both in terms of the timings of the annual peaks and in the integrated annual flux. Interannual variations in the integrated flux are of similar magnitude for both the model output and sediment trap measurements, but there is no significant relationship between these two sets of estimates. No long-term trend in flux is evident, either from the model, or from the measurements.During two spring/summer periods, the marine snow concentration in the water column was assessed by time-lapse photography and showed a strong peak at the start of the downward pulse of material at 3000 m. This emphasises the importance of large particles during periods of maximum flux and at the start of flux peaks. Time lapse photographs of the seabed show a seasonal cycle of coverage of phytodetrital material, in agreement with the model output both in terms of timing and magnitude of coverage prior to 1996. However, after a change in the structure of the benthic community in 1996 no phytodetritus was evident on the seabed.The model output shows only a single peak in flux each year, whereas the measured data usually indicated a double peak. It is concluded that the observed double peak may be a reflection of lowered sediment trap efficiency when flux is very high and is dominated by large marine snow particles.Resuspension into the trap 100 m above the seabed, when compared to the primary flux at 3000 m depth (1800 mab) was lower during periods of high primary flux probably because of a reduction in the height of resuspension when the material is fresh. At 2 mab, the picture is more complex with resuspension being enhanced during the periods of higher flux in 1997, which is consistent with this hypothesis. However there was rather little relationship to flux at 3000 m in 1998.At 3000 m depth, the Flux Stability Index (FSI), which provides a measure of the constancy of the seasonal cycle of flux, exhibited an inverse relationship with flux, such that the highest flux of organic carbon was recorded during the year with the greatest seasonal variation.     

Nutrient variability during El Niño 1997–98 in the California current system off central California

Nutrient conditions off central California during the 1997–98 El Niño are described. Data were collected on 11 cruises from March 1997 to January 1999 along a hydrographic section off central California, as well as every two weeks at a coastal station in Monterey Bay. Perturbations associated with El Niño are shown as anomalies of thermohaline and nutrient distributions along this section. The anomalies were obtained by subtracting seasonal averages for the period from April 1988 to April 1991 from the 1997–98 observations. The first indications of El Niño conditions (high sea levels) were observed at Monterey between late May and early June 1997, but the coastal nutricline did not begin to deepen until August 1997. It reached maximum depth of 130 dbar in January 1998 at the time that maximum sea level anomalies were observed. During this period: (1) the highest subsurface temperature anomalies coincided with subsurface nutrient anomaly minima at the depth of the pycnocline; (2) southern saline and nutrient-poor waters occupied the upper 80 dbar of the water column along the entire section; and (3) nitrate levels were close to zero in the euphotic zone, collapsing the potential new primary production in the coastal domain. At the end of February 1998, the nutricline shoaled to 40 dbar at the coast although it remained anomalously deep offshore. Higher temperatures and lower nutrient levels were observed for the entire section through August 1998 although in contrast with the previous winter, there was a strong freshening mainly due to an onshore movement of subarctic waters.     

Contribution of benthic microalgae to the whole water algal biomass and primary production in Suo Nada,the Seto Inland Sea,Japan

Biomass and primary productivity of benthic microalgae (BMA) and planktonic algae in Suo Nada, the western part of the Seto Inland Sea, Japan were compared in terms of unit area with regard to their seasonal and spatial distribution in 2002. Judging from light compensation depth and water depth, the southwestern part of Suo Nada was considered to be a potential habitat for BMA. Whereas the contribution of sedimented planktonic algae was high in biomass at the sediment surface, BMA was obviously significant both in biomass and primary production in the shallow southwestern part. However, the contribution of BMA to the total biomass in the entire water column was 7% in winter and 2% in summer. The primary production of BMA varied between 4.0 and 74.0 mg C m⁻² d⁻¹ in the southwestern part, accounting for 2–12% of the whole water column primary production. The ecological roles of BMA in the Suo Nada ecosystem are discussed, such as reduction of benthic nutrient flux, oxidation of surface sediments and feed for higher animals.     

Particle dynamics in the deep water column of Sagami Bay, Japan. I: origins of apparent flux of sinking particles

Temporal variations of sinking particle flux, together with their organic chemical properties, were monitored in the deep basin of Sagami Bay, Japan, using sediment traps with very high time resolutions from March 1997 to August 1998. At a height of 350 m above the bottom (about 1200 m water depth), the averaged total mass flux was more than 1000 mg/m²/day, which is about 10 times higher than those obtained for open ocean regions near Sagami Bay. While large amounts of phytodetritus, derived from phytoplankton blooms in the surface water, were transported downward in spring, the following extraordinary patterns in the temporal variability of sinking particle flux were also observed: (1) A sustained large flux of sinking particles during low productive periods from summer to winter in 1997. (2) An episodic increase of sinking particle flux in June 1998. (3) A difference in the temporal variability of sinking particles between the spring bloom periods of 1997 and 1998. The content of total organic carbon (TOC) and the stable carbon isotopic ratio (δ¹³C) of TOC demonstrated that the large fluxes observed in (1) and (2) could be attributed to the resuspension of phytodetritus deposited on the sea floor during the spring bloom period, and the abrupt erosion of surface sediment on the continental slope, respectively. The concentration of suspended particles in the deep water column affect the apparent flux of sinking particles. At the same time, sinking particles exported from surface waters during the spring bloom both decrease and increase suspended particle concentration through scavenging and rebound processes, respectively. Finally, the apparent difference in sinking particle flux between 1997 and 1998, (3), could be explained by differences in the extent of the scavenging process, which depend on the flux and quality of exported particles from the surface waters.     

Impact of the 1997–1998 El Niño and 1999 La Niña on nutrient supply in the Gulf of Alaska

Nutrient surveys of the Gulf of Alaska, from 1997 through 1999, show that coastal waters of British Columbia and southern Alaska experienced nitrate depletion each spring and summer. Through the 1997–1998 El Niño, waters with less than 1 μM NO₃ covered 250,000 km² area greater than 1999. Silicate levels as low as 0.2 μM were observed in coastal waters, suggesting that diatom growth may have been nutrient limited both in 1998 and 1999. Detailed sampling off the southern coast of British Columbia revealed that 1998 nitrate levels were only half the average of that during the 1970s winter, were depleted 1 month earlier in spring and remained low throughout the summer. Satellite images show that, compared to 1997 and 1999, chlorophyll levels were much lower in the spring of 1998 throughout the coastal waters of the Gulf of Alaska. Conditions changed dramatically during the 1999 La Niña, with ocean-mixed layer depths increasing by 20 m in winter and 40 m in spring when compared to that during 1997–1998 El Niño. Winter nutrient levels increased and summer upwelling returned. Over the past several decades, a trend towards greater stratification of coastal waters appears to be affecting the supply of nutrients to the mixed layer. The effects of stratification were especially obvious during the 1998 El Niño.     

Temporal variation in phytoplankton composition related to water mass properties in the central part of Sagami Bay

Temporal variations in water mass properties and the composition of phytoplankton pigments in the central part of Sagami Bay were investigated by monthly observations from June 2002 to May 2004. Eleven pigments were quantified using high-performance liquid chromatography (HPLC) from 100%, 20%, and 5% light depths relative to the surface; the class-specific composition of phytoplankton community was then obtained by CHEMTAX analysis. The study area was influenced by the Kuroshio water for most of the observation period. The mean contribution of diatoms in all samples was relatively low (29%), while that of flagellates, mainly chlorophytes or cryptophytes, was quite high (60%). The phytoplankton composition at the three depths was uniform throughout the observation period, indicating that the vertical structure of the phytoplankton community did not develop significantly over time. A distinct temporal pattern was observed: flagellates dominated during the summer of 2002 and the winters of 2002–2003 and 2003–2004, while diatoms dominated during the summer of 2003. This pattern was associated with water mass changes. The community in the summer of 2003 was influenced by coastal water. While no distinct spring bloom of phytoplankton was observed, a weak increase in chlorophyll a was observed during the spring of 2004. Ocean color satellite data showed that fluctuations in chlorophyll a concentrations at time scales much shorter than a month occurred during the spring of 2003 and that the elevations in chlorophyll a levels were not continuous. The fluctuations were probably associated with rapid flushing by the Kuroshio water, which has low chlorophyll a content.     

Benthic Disturbance and Impact Experiments in the Central Indian Ocean Basin

As a part of the Environmental Impact Assessment studies for nodule mining, a long-term program has been initiated in the Central Indian Basin. Multidisciplinary studies on geological, biological, physical, and chemical parameters were carried out in an area selected on the basis of baseline data collected in the first phase of the program. A benthic disturbance was simulated with a hydraulic device also used in the previous experiments in the Pacific Ocean. A site of 3,000 ×200 m was repeatedly disturbed by a combination of fluidizing pump and suction pump to dislodge and discharge sediment from the seafloor into the water column 5 m above the seafloor. During 9 days of operation, 26 tows were carried out for 47 h of disturbance, resuspending about 6,000 m 3 of sediment along an 88-km line. Data for postdisturbance impact assessment were collected with sediment traps, deep-towed cameras, seafloor samples, and conductivity-temperature depth sensor (CTD)-rosette observations. Seafloor data, sediment samples, and water column studies were aimed at evaluating the impact of benthic disturbance, on the basis of pre- and postdisturbance data collected during the experiment. Observations show that vertical mixing of sediment as well as its lateral movement and resedimentation because of plume migration alters various parameters and leads to changes in the environment around the area.     

Has the 1998 regime shift also occurred in the oceanographic conditions and lower trophic ecosystem of the Oyashio region?

To examine whether the regime shift in 1998 that has been variously reported to have occurred in the oceanographic conditions of the central and eastern North Pacific also occurred in the Oyashio region, western North Pacific, we compared data over the period 1990–2003. Oceanographic conditions were compared before 1997 with those after 1998, using the A-line dataset (1990–2003) obtained by the oceanographic surveys of the Hokkaido National Fisheries Research Institute, Fisheries Research Agency (HNFRI/FRA). Seasonal changes of the monthly-mean SST (as temperature in the surface layer) show a significant increase in spring after 1998. After 1998, the mean concentration of chlorophyll a at the surface was higher in spring than that before 1997. This was more remarkable in the main current of the Oyashio. These changes suggest that the spring phytoplankton bloom in the Oyashio region after 1998 was larger in magnitude and initiated earlier. Consumption of nutrients during the spring bloom and standing stock of netplankton also shows a distinct difference between the time period before 1997 and after 1998. These results support the occurrence of the regime shift around 1998 in the Oyashio region. The changes of hydrographical conditions accompanying with the 1998 regime shift are discussed. The hydrographic mechanism of enhancement of primary productivity during the spring phytoplankton bloom was not fully clarified, though. Results in this study may support the usefulness of the A-line dataset for analysis of long-term variability in the western subarctic Pacific.     

The PROWQM physical–biological model with benthic–pelagic coupling applied to the northern North Sea

PROWQM, a 1-D depth resolving model which couples physical and microbiological processes in the water column with sedimentation/resuspension and benthic mineralisation processes, has been used to simulate seasonal changes of chlorophyll, nutrients and oxygen at the PROVESS north site (59°20′N 1°00′E) in the North Sea. PROWQM is derived from the 3-D model COHERENS, and improves COHEREN's benthic and pelagic biology.The physical sub-model of PROWQM implicitly solves turbulence closure equations forced by climatological, or realistic high-frequency, meteorological and tidal data. The pelagic biological sub-model 2MPPD includes a ‘diatomy’ microplankton (mp1) and a ‘flagellatey’ (or microbial loop) microplankton (mp2), the cycling of silicon and nitrogen, slow-sinking detritus, and fast-sinking phytodetritus. Phytodetritus is formed by shear-driven aggregation of particulate material, using a simple algorithm for bulk processes that is derived by considering the interactions of single cells. The microplankton compartments include heterotrophic bacteria and protozoa as well as phytoplankton, and most microplankton rates are specified with the aid of a ‘heterotroph fraction’ parameter, which was 0.125 for mp1 and 0.6 for mp2. The microbiological system is closed by mesozooplankton grazing pressures imposed as time varying series determined from observed zooplankton abundance. The benthic boundary sub-model includes a superficial fluff layer and a nutrient element reservoir in the consolidated sediment. Particulate material in the fluff layer can be resuspended (in response to bed stress by near-bed flows), mineralised or carried by bioturbation into the underlying, consolidated, sediment, where it is mineralised and its nutrients returned to the water-column at rates mainly dependent on (implicit) macrobenthic pumping. Benthic denitrification can occur when mineralisation rates exceed oxygen supply.Verification of the PROWQM numerical implementation used test cases and checks for nutrient element conservation. Simulations with realistic forcing, for a range of parameter values, were compared with historic observations in the NOWESP data set and during FLEX76, and with those made during the PROVESS cruises in autumn 1998. PROWQM provided a good simulation of the seasonal succession from a diatom-dominated spring bloom to summer dominance by small flagellates. The simulations included sedimentation of organic matter from the spring bloom, and qualitatively realistic behaviour of the fluff layer, but decay rates were too slow and there was almost no denitrification. The simulated surface mixed layer was too shallow during the summer. Simulated annual net microplankton primary production was in between 59 and 91 g C m⁻² y⁻¹. A large proportion of mineralisation, 28–47% of nitrogen and 40–67% of silicon mineralisation, took place as a result of the decay of sinking and resuspended detritus whilst in the water column.PROWQM is discussed in relation to other models that have been used to simulate this part of the North Sea, in particular the simpler ECOHAM1 and the more complex ERSEM, and in relation to PROWQM's evolution from COHERENS.     

Copepod grazing on phytoplankton in the Pacific sector of the Antarctic Polar Front

Mesozooplankton abundance, community structure and copepod grazing on phytoplankton were examined during the austral spring 1997 and summer 1998 as part of the US JGOFS project in the Pacific sector of the Antarctic polar front. Mesozooplankton abundance and biomass were highest at the polar front and south of the front. Biomass increased by 1.5–2-times during the course of the study. Calanoides acutus, Calanus propinquus, C. simillimus, Rhincalanus gigas and Neocalanus tonsus were the dominant large copepods found in the study. Oithona spp and pteropods were numerically important components of the zooplankton community. The copepod and juvenile krill community consumed 1–7% of the daily chlorophyll standing stock, equivalent to 3–21% of the daily phytoplankton production. There was an increased grazing pressure at night due to both increased gut pigment concentrations as well as increases in zooplankton numbers. Phytoplankton carbon contributed a significant fraction (>50%) of the dietary carbon for the copepods during spring and summer. The relative importance of phytoplankton carbon to the diet increased south of the polar front, suggested that grazing by copepods could be important to organic carbon and biogenic silica flux south of the polar front.     

泉州湾水域浮游植物群落的昼夜变化

对福建省泉州湾口、大坠岛以北水域的浮游植物群落作了准同步昼夜连续观测,观测时间分别在2001年11月至2002年8月内的秋(11月)、冬(2月)、春(5月)、夏(8月)季大潮期.结果表明,该水域的叶绿素a生物量(以下简称生物量)受湾内高生物量的影响在潮汐作用下出现波动,高潮期出现低生物量,低潮期出现高生物量.靠近内湾的站位生物量大于湾外的站位;底层水体中的生物量普遍大于表层,底栖硅藻成为生物量变化的重要部分.调查海区生物量呈现昼夜节律变化,生物量白天高于夜间.进一步的分析表明,在潮汐和昼夜变化综合作用下,浮游植物的群落结构发生相应变化,白天的多样性指数低于夜间,浮游植物昼夜群落的不相似度很高.群落中一些种类如中肋骨条藻白天大量增殖,出现较高的优势度,昼夜平均密度比值达到9,变化节律明显.研究认为,尽管海区潮汐和流向左右着生物量变化,浮游植物的生态学过程如细胞增殖、再悬浮、扩散等过程在昼夜生物量变化中仍产生较大作用.     

Benthic mineralization rates at two locations in the southern North Sea

Benthic oxygen uptake, sulphate reduction and benthic bacterial production were measured at two contrasting locations in the southern North Sea: the shallow and turbulent Broad Fourteens area in the Southern Bight, and the deeper Oyster Grounds, a deposition area, where thermohaline stratification occurs during summer. Oxygen uptake and sulphate reduction showed a clear seasonal pattern in the Broad Fourteens area, indicating a supply of carbon to the benthic system that is closely related to the standing stock of carbon in the water column. This close benthic-pelagic coupling is probably due to the influence of the tide in this part of the North Sea, which keeps the water column permanently mixed. At the Oyster Grounds, no seasonal pattern was observed. Peaks in oxygen uptake and sulphate reduction were found in winter. Irregularly occurring events, such as storms and fishery-related activities, are likely to affect the benthic mineralization patterns in this area. Annual benthic carbon mineralization rates estimated from oxygen uptake rates were 44 gC·m⁻² at the Broad Fourteens, and 131 gC·m⁻² at the Oyster Grounds, of which 26 and 28%, respectively, could be attributed to sulphate reduction (assuming an annual sulphide reoxidation rate of 100%). Although sulphate reduction rates in the southern North Sea are higher than previously suggested, aerobic respiration is the most important pathway for benthic carbon mineralization at the stations visited. Production rates of benthic bacterial carbon measured with labelled leucine were much higher than carbon mineralization rates based on oxygen uptake or sulphate reduction. This may either imply a very high bacterial carbon conversion efficiency, or point to shortcomings in the accuracy of the techniques. A critical evaluation of the techniques is recommended.     

Temporal variability of near-bottom particle resuspension and dynamics at the Porcupine Abyssal Plain, Northeast Atlantic

In order to study particle behaviour and its time-variability in the near-bottom layer on the Porcupine Abyssal Plain (48°50′N, 16°30′W, 4850 m), long-term measurements were made of currents, and nephelometry and particle samples were collected using an autonomous lander between mid-1996 and mid-1998. Water samples, collected in the Bottom Nepheloid Layer within 1000 m of the bottom, were filtered for suspended particles whose contents of organic carbon, nitrogen and pigments were determined. This study was co-ordinated with a water column flux study and a detailed programme of benthic studies to understand how the abyssal boundary layer responds to and modifies inputs of organic matter from the water column (MAST3/BENGAL programme).There were strong seasonal fluctuations in the near-bottom (2 m above the bottom, mab) particle flux, whose variation were correlated in time with the water column fluxes. During the periods of peak flux, the near-bottom flux was sometimes higher than that recorded higher up in the water column, but not always at other times. These excesses were attributed to the resuspension events, since we observed a correlation between current speed and nephelometry. However, in summer the peak in the particle resuspension flux could not be explained by the variations in the tidal amplitude. Instead we attribute it to the large quantities of fresh large particles (aggregations) that had just arrived on the bottom; it was probably linked to the feeding activity and sediment reworking by the rich and varied benthic and benthopelagic megafauna.In both 1997 and 1998, the nephelometry signal (directly related to fine particle concentration) and its variability increased after the peaks in large particle flux with a time-lag of 2–3 months. We assume that this time lag corresponds to the time it takes for the large fresh particles, once they have settled on the bottom, to be disaggregated into smaller particles, and hence become subject to resuspension in the quiet current conditions then prevailing in the BENGAL area. The suspended particle analyses confirm the vertical structure of the Bottom Nepheloid Layer, the lower part of which corresponds to the Bottom Mixed Layer (BML) where resuspension and mixing are higher.     

Effects of the 1997–1999 El Niño and La Niña events on zooplankton abundance and euphausiid community composition within the Monterey Bay coastal upwelling system

Zooplankton abundance and euphausiid community composition were sampled seasonally (spring, summer, fall) within Monterey Bay, California, between 1997 and 1999. Measurements of sea surface temperature (SST), mixed layer depth, and upwelling indices provided concurrent data on physical oceanographic parameters. Both total zooplankton and krill abundance dramatically declined in the summer of 1997 coincident with a rapid increase in SST and mixed layer depth. Changes in euphausiid community composition occurred in concert with the decline in overall abundance. The relative abundance of the southern neritic Nyctiphanes simplex increased from August to November in 1997, the abundance of cold temperate Euphausia pacifica decreased significantly, and that of the northern neritic Thysanoessa spinifera declined dramatically. The sudden appearance of an adult cohort of N. simplex in July 1997 suggests that rapid poleward flow characteristic of coastally trapped Kelvin waves occurred between June and July of 1997. The persistent presence of warm temperate and subtropical taxa in samples collected between August 1997 and October 1998 indicates that this poleward flow continued in 1998. Zooplankton abundance, euphausiid community composition, and physical oceanographic parameters gradually returned to a more typical upwelling-dominated state in the spring and summer of 1998. E. pacifica and T. spinifera abundances gradually increased during the summer and fall of 1998, while N. simplex abundance abruptly declined in the spring of 1998. However, this recovery was confined to a narrow coastal band as a result of the onshore movement of the oceanic waters of the California Current. This was reflected by higher than normal numbers of the oceanic Nematoscelis difficilis within samples collected during the spring and summer of 1998. By the spring and summer of 1999, both zooplankton and euphausiid abundance had increased to the highest levels recorded during the 3-year study. Both E. pacifica and T. spinifera abundance increased relative to 1998 while N. simplex was completely absent in all samples. These changes reflected the cooler, highly productive environmental conditions associated with the 1998/1999 La Niña.     

Life Cycle,Growth and Secondary Production in a Brackish-Water Population of the Polychaete Notomastus latericeus (Capitellidae) in the Mediterranean Sea

Abstract. Three years of observations on a population of Notomastus latericeus Sars from Acquatina lagoon (Lecce, Italy) are reported. The population dynamics, life cycle, and reproduction were investigated in order to explain periodic density variations of this species. Some physical parameters of the water column were measured and sediment analyses were performed to characterize the environment. Secondary production was also estimated as part of a functional study of the benthic system in the Acquatina lagoon. The species is opportunistic; its density is apparently correlated to variation in salinity, but its presence in the lagoon is dependent on recruitment success and competition with other polychaete species.     

Recovery of the macrobenthic community in the Valli di Comacchio,northern Adriatic Sea,Italy

The macrobenthic community structure of the Valli di Comacchio (northern Italy) was described in order to assess the ecological conditions of the main basins through 2001. In addition, the biotic data set gathered in 1997–1998 was compared to achieve an estimation of the eventual recovery of the lagoonal benthic assemblages. Four permanent sites (P1, M4, M5 and M6), located along a gradient of sediment texture and confinement, and representing four different areas of the lagoon, were sampled quarterly in 2001 for macrofauna and sedimentary variables (organic content, phytobenthic chlorophyll-a, depth of the redox potential discontinuity layer), and fortnightly for water variables (temperature, salinity, dissolved oxygen and planktonic chlorophyll-a). Water and sedimentary variables were typical of eutrophic areas. A total of 52 macrobenthic taxa were identified and, on the basis of the species collected, differences in faunal distribution among the four areas were recognized, although less marked than in previous studies. Sediment trophic status and its seasonal dynamics in the Valli were crucial in determining species distribution among the different areas. The comparison between 1997–1998 and 2001 biotic data indicated that conditions in the lagoon had improved, especially in the formerly most impaired, central area. In the Valli di Comacchio, the recovery of benthic communities after severe disturbance will probably take longer, even if sewage discharges have been removed 11 years ago. Secondary disturbance due to scarce water circulation, sharp temperature and salinity fluctuations, release of toxic substances from sediments influenced animal assemblages along spatial and temporal scales. The interplay of these variables probably caused deviations from the expected improvement in benthic conditions. Nevertheless, clear signs of amelioration, particularly at the previously most impaired area, were detectable.     

Trophic efficiency of the planktonic food web in a coastal ecosystem dominated by Phaeocystis colonies

The trophic efficiency of the planktonic food web in the Phaeocystis-dominated ecosystem of the Belgian coastal waters was inferred from the analysis of the carbon flow network of the planktonic system subdivided into its different trophodynamic groups. A carbon budget was constructed on the basis of process-level field experiments conducted during the spring bloom period of 1998. Biomass and major metabolic activities of auto- and heterotrophic planktonic communities (primary production, bacterial production, nanoproto-, micro- and mesozooplankton feeding activities) were determined in nine field assemblages collected during spring at reference station 330. In 1998, the phytoplankton spring flowering was characterised by a moderate diatom bloom followed by a massive Phaeocystis colony bloom. Phaeocystis colonies, contributing 70% to the net primary production, escaped the linear food chain while the early spring diatom production supplied 74% of the mesozooplankton carbon uptake. The rest of mesozooplankton food requirement was, at the time of the Phaeocystis colony bloom, partially fulfilled by microzooplankton. Only one-third of the microzooplankton production, however, was controlled by mesozooplankton grazing pressure. Ungrazed Phaeocystis colonies were stimulating the establishment of a very active microbial network. On the one hand, the release of free-living cells from ungrazed colonies has been shown to stimulate the growth of microzooplankton, which was controlling 97% of the nanophytoplankton production. On the other hand, the disruption of ungrazed Phaeocystis colonies supplied the water column with large amounts of dissolved organic matter available for planktonic bacteria. The budget calculation suggests that ungrazed colonies contributed up to 60% to the bacterial carbon demand, while alternative sources (exudation, zooplankton egestion and lysis of other organisms) provided some 30% of bacterial carbon requirements. This suggests that the spring carbon demand of planktonic bacteria was satisfied largely by autogenic production. The trophic efficiency was defined as the ratio between mesozooplankton grazing on a given source and food production. In spite of its major contribution to mesozooplankton feeding, the trophic efficiency of the linear food chain, restricted to the grazing on diatoms, represented only 5.6% of the available net primary production. The trophic efficiency of the microbial food chain, the ratio between mesozooplankton grazing on microzooplankton and the resource inflow (the bacterial carbon demand plus the nanophytoplankton production) amounted to only 1.6%. These low trophic efficiencies together with the potential contribution of ungrazed Phaeocystis-derived production to the bacterial carbon demand suggest that during spring 1998 most of the Phaeocystis-derived production in the Belgian coastal area was remineralised in the water column.     

Anomalous hydrographic and biological conditions in the northern South China Sea during the 1997–1998 El Niño and comparisons with the equatorial Pacific

It is demonstrated that weakened wind mixing and strengthened water column stratification resulted in the anomalously low sea surface chlorophyll in the northern South China Sea during the 1997–1998 El Niño event. Remotely sensed sea surface temperature, wind and chlorophyll, which were validated by shipboard observations at the SouthEast Asian Time-series Study (SEATS) station (18°N, 116°E) in the northern South China Sea (SCS) provided the basis for this study. During the 1997–1998 winter at the SEATS station, the sea surface temperature was elevated by about 2 °C above the climatological mean, while the wind speed of the northeast monsoon was reduced from a climatological mean of 9.4 to 6.8 m/s. The concentration of surface chlorophyll-a dropped from 0.2 to 0.1 mg/m³. The monthly area-averaged integrated primary production estimated for the northern SCS area (112–119°E, 15–21°N) was reduced by about 40% of the normal winter value. Under the anomalously high sea surface temperature and weak monsoon, the mixed-layer depth would have been reduced from an average of 65 to 45 m and the nutrients in the mixed layer would have been reduced by half, according to observations at the SEATS station in more recent years. During the 1997–1998 El Niño event, the onset of warming in the northern SCS lagged behind that in the eastern equatorial Pacific by about 5 months and lingered for 11 months. This course of change resembled that of the western Pacific warm pool region. However, contrary to the northern SCS, the sea surface chlorophyll was enhanced in the warm pool region during the event, probably mainly because of the uplifted nutricline. Unlike the eastern equatorial Pacific, the dramatic recovery of biological production did not happen in the SCS in the summer of 1998. These distinctive biogeochemical responses reflect fundamental differences between the SCS and the equatorial Pacific in terms of upper water column dynamics.