Seasonal and inter-annual variation of mesozooplankton in the coastal upwelling zone off central-southern Chile

Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency of 30 days (August 2002 to December 2005), allowed the assessment of seasonal and inter-annual variation in zooplankton biomass, its C and N content, and the community structure in relation to upwelling variability. Copepods contributed 79% of the total zooplankton community and were mostly represented by Paracalanus parvus, Oithona similis, Oithona nana, Calanus chilensis, and Rhincalanus nasutus. Other copepod species, euphausiids (mainly Euphausia mucronata), gelatinous zooplankton, and crustacean larvae comprised the rest of the community. Changes in the depth of the upper boundary of the oxygen minimum zone indicated the strongly seasonal upwelling pattern. The bulk of zooplankton biomass and total copepod abundance were both strongly and positively associated with a shallow (<20 m) oxygen minimum zone; these values increased in spring/summer, when upwelling prevailed. Gelatinous zooplankton showed positive abundance anomalies in the spring and winter, whereas euphausiids had no seasonal pattern and a positive anomaly in the fall. The C content and the C/N ratio of zooplankton biomass significantly increased during the spring when chlorophyll-a was high (>5 mg m⁻³). No major changes in zooplankton biomass and species were found from one year to the next. We concluded that upwelling is the key process modulating variability in zooplankton biomass and its community structure in this zone. The spring/summer increase in zooplankton may be largely the result of the aggregation of dominant copepods within the upwelling region; these may reproduce throughout the year, increasing their C content and C/N ratios given high diatom concentrations.     

Fine-scale vertical distribution of coastal and offshore copepods in the Golfo de Arauco, central Chile, during the upwelling season

In order to understand the mechanism by which zooplankters from different origins co-occur during the upwelling season within Golfo de Arauco, one of the most productive areas in central Chile, we assessed short term variations in the vertical distribution of the most abundant copepod species. Fine-scale, day and night vertical zooplankton sampling was done with a pump over 12 days in summer. The water column in the gulf consisted of three layers: Equatorial Subsurface Water of low dissolved oxygen content in the deeper part of the water column, strong temperature and oxygen gradients at mid-depth (15-25 m), and a layer of warmer, more oxygenated, less saline water at the surface. Copepods within the gulf originated from offshore, from the continental shelf, and from the coastal area. Most taxa showed distinctive vertical distributions. Three copepod groups were identified by their mean weighted depths of residence. One group included shallow residents found above the thermocline/oxycline (Acartia tonsa, Centropages brachiatus, Corycaeus sp., Paracalanus parvus, Oncaea sp.). A second group was comprised by species distributed at or below the thermocline/oxycline (Oithona sp., Oncaea conifera, Lucicutia sp., Metridia sp., Heterorhabdus papilliger). The third group was composed of vertical migrators that crossed the thermocline/oxycline (Calanus chilensis, Calanoides patagoniensis, Aetideus armatus, Pleuromamma piseki). In spite of their different vertical distribution ranges, the most abundant and frequent copepod species (P. parvus, C. chilensis, C. patagoniensis, C. brachiatus) share a common capacity to withstand wide ranges of oxygen concentration and temperature. This characteristic, along with the capacity to vary their life strategies under different environmental conditions, seems to facilitate the maintenance of large numbers of copepods in coastal waters along the Humboldt Current.     

Cross-shelf distribution of copepods and the role of event-scale winds in a northern California upwelling zone

Cross-shelf distribution and abundance of copepod nauplii and copepodids were measured during three summer upwelling seasons (2000–2002) in a coastal upwelling zone off northern California. These 3 years varied considerably in the intensity of winds, abundance of chlorophyll, and water temperature. The cruises in 2000 were characterized by relaxation conditions, with generally high levels of chlorophyll and high water temperature. The cruises in 2001 and 2002 were dominated by strong and persistent upwelling events, leading to lower chlorophyll and water temperatures. The copepod assemblage was dominated by Oithona spp., Acartia spp. and Pseudocalanus spp., with Metridia pacifica (lucens), Microsetella rosea, Oncaea spp. and Tortanus discaudatus also common during all 3 years. The cross-shelf distribution of copepods was generally shifted offshore during upwelling and onshore during relaxation events, although some variability between species occurred. Abundance of all life stages generally exhibited a negative correlation with cross-shelf transport averaged over at least 1–4 days and lagged by 0–3 days, indicating lower abundances during and immediately after active upwelling. However, copepod nauplii seemed to respond positively to wind events lasting 1–5 days followed by a period of relaxation lasting 6 or 7 days. These rapid rates of change in abundance are probably too great to be due to in situ growth and reproduction alone; physical processes must also play a role. These results suggest a highly dynamic relationship between copepods and upwelling events off northern California, with species-specific responses to upwelling to be expected.     

Biogeography and phenology of satellite-measured phytoplankton seasonality in the California current

Thirteen years (1998–2010) of satellite-measured chlorophyll a are used to establish spatial patterns in climatological phytoplankton biomass seasonality across the California Current System (CCS) and its interannual variability. Multivariate clustering based on the shape of the local climatological seasonal cycle divides the study area into four groups: two with spring-summer maxima representing the northern and southern coastal upwelling zones, one with a summer minimum offshore in mid-latitudes and a fourth with very weak seasonality in between. Multivariate clustering on the seasonal cycles from all 13 years produces the same four seasonal cycle types and provides a view of the interannual variability in seasonal biogeography. Over the study period these seasonal cycles generally appear in similar locations as the climatological clusters. However, considerable interannual variability in the geography of the seasonal cycles is evident across the CCS, the most spatially extensive of which are associated with the 1997–1999 El Niño-Southern Oscillation (ENSO) signal and the 2005 delayed spring transition off the Oregon and northern and central California coasts. We quantify linear trends over the study period in the seasonal timing of the two seasonal cycles that represent the biologically productive coastal upwelling zones using four different metrics of phenology. In the northern upwelling region, the date of the spring maximum is delaying (1.34 days yr⁻¹) and the central tendency of the summer elevated chlorophyll period is advancing (0.63 days yr⁻¹). In the southern coastal upwelling region, both the initiation and cessation of the spring maximum are delaying (1.78 days yr⁻¹ and 2.44 days yr⁻¹, respectively) and the peak is increasing in duration over the study period. Connections between observed interannual shifts in phytoplankton seasonality and physical forcing, expressed as either basin-scale climate signals or local forcing, show phytoplankton seasonality in the CCS to be influenced by changes in the seasonality of the wind mixing power offshore, coastal upwelling in the near-shore regions and basin-scale signals such as ENSO across the study area.     

Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

Seasonal dynamics of sub-ice fauna below pack ice in the Arctic (Fram Strait)

Pronounced seasonality is a characteristic feature of polar ecosystems, but seasonal studies in the high-Arctic pack-ice zone are still scarce because of logistical constraints. During six expeditions (1994–2003) to the Fram Strait area between Greenland and Svalbard in winter, spring, early summer, late summer and autumn, the sub-ice habitat and fauna below the pack ice (0–1 m depth) were analyzed for seasonal patterns. Both environmental variables such as ice cover, temperature, salinity and chlorophyll a (chl a), as well as species composition, abundance and biomass of the sub-ice fauna showed distinct seasonal dynamics. Most species of the sub-ice fauna were found in early summer, followed by autumn, spring and late summer; the lowest number occurred in winter. The sub-ice fauna was dominated by copepod nauplii during all seasons. Next numerous was the small pelagic copepod Oithona similis, followed by occassional swarms of Pseudocalanus minutus and Calanus spp. Abundances of the sympagic fauna in the sub-ice water layer were much lower, with ectinosomatid copepods being usually the most numerous sympagic group. In the course of the year, total abundances of the sub-ice fauna showed a steep increase from the earliest sampling dates towards the end of winter/beginning of spring reaching maximum numbers then, and a decrease to minimum numbers in early summer. A second peak occurred in late summer, followed by a decrease towards autumn. This significant trend was due to the abundances of copepod nauplii and Oithona similis. Sympagic species were virtually absent during winter, and increased significantly in spring and early and late summer. A factor analysis revealed the variables ice cover and thickness, water temperature and salinity, as well as chl a as the major controlling factors for the seasonal patterns in different groups and species of the sub-ice fauna. Because of the special environmental conditions in the sub-ice habitat, and the unique species composition characterized by small taxa, young stages, and sympagic species, the seasonal dynamics of the Arctic sub-ice fauna differ substantially from those of the epipelagic zooplankton community in the Arctic Ocean.     

Mesozooplankton community characteristics in the NE subarctic Pacific

Mesozooplankton biomass, species composition, abundance, and vertical distribution were determined along a transect from the continental slope off the west coast of Canada to Ocean Station Papa (OSP) in the open-ocean waters of the NE subarctic Pacific as part of the Canadian Joint Global Ocean Flux Study of this area. All of these measurements had distinct seasonal patterns. At OSP biomass peaked in spring, coincident with the annual biomass maximum of large copepods of the genus Neocalanus. Early copepodites of these copepods were present in surface waters at all stations along the transect in winter, but N. plumchrus and N. flemingeri copepodites were only at the offshore stations in spring. This indicated that these large copepods had completed the growth phase of their life cycle slower in the open ocean than closer to shore where they had already descended to deep water by May or June. Summer biomass was low compared to the spring peak. The summer mesozooplankton abundance was similar to the springtime abundance, but the composition had changed from large-bodied copepods in the spring to small copepods and fewer non-copepod taxa in the summer, which accounts for the reduction in total biomass. Winter biomass was the lowest of the year. Winter species composition was similar to summer except for the appearance of juvenile stages of the genera Neocalanus and Calanus. Diel changes in biomass in the upper 150 m were found in summer but not in winter or spring. Vertical distributions of copepods were often distinct, with closely related species occupying different depth strata. Measurements of wet weight at OSP were higher than the long-term mean wet weight during winter and spring, and lower during summer.     

Population succession and feeding of scyphomedusae, Aurelia aurita, in a eutrophic tropical lagoon in Taiwan

Seasonal dynamics and feeding of scyphomedusae, Aurelia aurita, were investigated monthly from 1999 to 2002 in relation to environmental conditions in Tapong Bay, a eutrophic tropical lagoon in southwestern Taiwan. Medusae appeared throughout the year but exhibited seasonal dynamics that were correlated with hydrographic features in the bay. Most ephyrae of A. aurita occurred mainly in the lower flushed and eutrophic inner bay, and during the cold, dry season between November and February. They grew to young medusae with a maximum abundance in spring (March–May), but their numbers abruptly decreased during the warm and rainy summer season in June–September. The remaining medusae then grew rapidly to a maximal size of 29 cm. Mature females spawned in the following autumn when precipitation decreased but zooplankton food was still abundant. These mature individuals decreased in size after spawning and in winter. Gut content analysis revealed that A. aurita fed mainly on copepods and copepod nauplii and less on bivalve larvae and fish eggs. Prey selectivity indices indicated that larger medusae selected positively for copepods while small size medusae preferred copepod nauplii. The overall feeding effect of A. aurita on the standing stock of zooplankton was significant (27%) in the bay. Our results suggest that tidal flow and dense oyster culture pens were the two most important factors influencing the spatial distribution pattern of A. aurita in the bay, while precipitation affected the population abundance seasonally; decreasing water temperature coincided with the mass release of ephyra in late autumn and winter.     

台湾以北海域浮游桡足类生物海洋学特征的研究Ⅰ.数量分布

根据１９８７～１９９０年间在台湾以北海域进行的６个航次的调查资料,对该海域浮游桡足类数量分布作了探讨,结果表明,本区浮游桡足类丰度在春季最高,冬季最低,夏、秋季介于中间;黑潮锋内侧的桡足类丰度一般高于黑潮锋外侧,较高丰度区的分布具夏季最向外海、冬季最靠近岸、其他两季居中的趋势,这与黑潮锋位的季节摆动趋势基本相一致;浮游桡足类丰度的较高值区一般出现在不同水系的交汇区,尤其是黑潮次表层水涌升域的近岸边缘侧,而在黑潮次表层水的主水体中,因溶解氧含量低等因素,其丰度一般均较低;浮游桡足类丰度垂直分布趋势为随水深增加而减低,生物本身昼夜垂直移动规律可能是局部区域产生异常分布的原因之一．     

Water-body preferences of dominant calanoid copepod species in the Angola-Benguela frontal zone

The distribution of five dominant calanoid copepods was related to different water masses in the Angola-Benguela Front system. Five water bodies were identified by principal component analysis, on the basis of abiotic parameter such as temperature, salinity, dissolved oxygen, phosphate, silicate, nitrate and nitrite. These parameters were reduced to single factors and arranged along two principal component axes. The copepod species incuded females and copepodites C5 of Calanoides carinatus and females of Metridia lucens, Centropages brachiatus, Nannocalanus minor and Aetideopsis carinata. The water bodies identified in the frontal system were related to currents, upwelling processes, an oxygen minimum layer and biological modification. The different copepod species, as well as the two ontogenetic stages of C. carinatus, showed clear preference for specific water bodies, and their behavioural and physiological adaptations to the environment are discussed.     

Latitudinal Distribution of CO2 Fugacity along 175°E in the North Pacific in 1992–1996

In order to examine latitudinal distribution and seasonal change of the surface oceanic fCO₂, we analyzed the data obtained in the North Pacific along 175°E during the NOPACCS cruises in spring and summer of 1992–1996. Except for around the equator where the fCO₂ was significantly affected by the upwelling of deep water, the latitudinal distribution of fCO₂ showed distinctive seasonal variation. In the spring, the fCO₂ decreased and then increased going southward with the minimum value of about 300 µatm around 35°N, while in the summer, the fCO₂ displayed high variability, showing minimum and maximum values at latitudes of around 44° and 35°N, respectively. It was also found that the fCO₂ was well correlated with the SST, but the relationship between the two was different for different hydrographic regions. In the subpolar gyre, the frontal regions between the Water-Mass Front and the Kuroshio bifurcation front, and between the Kuroshio bifurcation front and the Kuroshio Extension current, SST, DIC and TA influenced the seasonal fCO₂ change through seasonally-dependent biological activities and vertical mixing and stratification of seawater. In the central subtropical gyre and the North Equatorial current, the seasonal fCO₂ change was found to be produced basically by changes in SST and DIC. The summertime oceanic fCO₂ generally increased with time over the period covered by this study, but the increased rate was clearly higher than those expected from other measurements in the western North Pacific.     

Seasonal variation of the neustonic zooplankton community in southern waters of Korea

We investigated the seasonal variation of the neustonic zooplankton community in the surface layer (above 30 cm in depth) of 16 stations in southern waters of Korea from April 2002 to April 2003. In the neustonic community, copepods accounted for 59% of total organisms, as the most predominant group, while other dominant groups were invertebrate larvae, cladocerans, tunicates and fish eggs. Pontellid copepods, a major group of neustonic zooplankton, showed a marked seasonal variation and displayed a significant positive correlation with temperature (P < 0.05). Fish eggs appeared from spring to early summer in the inshore waters while tunicates and oceanic copepods (Eucalanus spp. and Euchaeta spp.) were dominant in the offshore waters during high temperature periods. This suggests that the distribution pattern of the neustonic zooplankton is seasonally affected by the hydrological conditions of in- and offshore waters. The abundance of zooplankton was higher in the surface layer (< 15 cm) than in the subsurface layer (15–30 cm), and was also higher at night than during the daytime. Of the dominant copepod families, the density of acartiid and pontellid copepods was higher in the surface layer than the subsurface layer. The density of acartiid and corycaeid copepods was significantly higher during daytime than nighttime. This indicates that the surface layer in temperate regions may provide a favorable habitat for neustonic copepods despite the dynamic environmental fluctuations.     

Zooplankton research off Peru: A review

A review of zooplankton studies conducted in Peruvian marine waters is given. After a short history of the development of zooplankton research off Peru, we review zooplankton methodology, taxonomy, biodiversity, spatial distribution, seasonal and interannual variability, trophodynamics, secondary production, and modelling. We review studies on several micro-, meso-, macro-, and meroplankton groups, and give a species list from both published and unpublished reports. Three regional zooplankton groups have been identified: (1) a continental shelf group dominated by Acartia tonsa and Centropages brachiatus; (2) a continental slope group characterized by siphonophores, bivalves, foraminifera and radiolaria; (3) and a species-rich oceanic group. The highest zooplankton abundances and biomasses were often found between 4–6°S and 14–16°S, where continental shelves are narrow. Species composition changes with distance from the shore. Species composition and biomass also vary strongly on short time scales due to advection, peaks of larval production, trophic interactions, and community succession. The relation of zooplankton to climatic variability (ENSO and multi-decadal) and fish stocks is discussed in the context of ecological regime shifts. An intermediate upwelling hypothesis is proposed, based on the negative effects of low upwelling intensity in summer or extremely strong and enduring winter upwelling on zooplankton abundance off Peru. According to this hypothesis, intermediate upwelling creates an optimal environmental window for zooplankton communities. Finally, we highlight important knowledge gaps that warrant attention in future.     

Copepod community structure during upwelling and non-upwelling seasons in coastal waters off Cochin, southwest coast of India

Vypeen, an island of Cochin estuarine system, acts as the spawning site of several marine and estuarine fishes. We assumed that, physical process(upwelling) make changes in hydrography and the production of chlorophyll a in coastal waters off Vypeen. These alterations can influence the zooplankton abundance and copepod community structure in that area. For justifying this hypothesis, samples were collected from Vypeen at 10 m and 30 m locations during January(pre-southwest monsoon), August(late-southwest monsoon) and November(postsouthwest monsoon) 2014. During August, subsurface water column was cool, nutrient rich and less oxygenated(signatures of upwelling) than in November and January. Maximum concentrations of nitrate and chlorophyll a were recorded during August. In the present study, 15 zooplankton groups were recorded; of which copepods were the most predominant group(73%–90%). Copepod density in the present study ranged between 527.2 ind./m~3 and 5 139.2 ind./m~3. Totally, 37 copepods species were reported in present study and copepod species richness was higher during January. The abundance of zooplankton and copepods was high in August during late-southwest monsoon. These variations were closely associated with the coastal upwelling in August and weakening of moderate upwelling in November. In SIMPER analysis, it was found that the copepods species distribution was similar within season and dissimilar between the three seasons. Moreover, the higher abundance of upwelling indicator species Temora turbinata was recorded in August, which confirms the signs of seasonal upwelling in Vypeen. The present study emphasized on the influence of hydrographical parameters associated with physical process, in governing the copepod community organization of the Vypeen Island.     

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.     

Relationships between water-column stratification,phytoplankton cell size and copepod fecundity in Long Island Sound and off central Chile

Changes in phytoplankton abundance and size brought on by changes in water-column stratification are shown to affect copepod fecundity. The response of copepods to changes in their food environment, as manifested by changes in rates of egg production, is compared for two different time-scales: seasonal-to-weekly in Long Island Sound, New York, and event-scale-to-daily off the coast of central Chile. The data sets discussed include a time-series of hydrographic and chlorophyll samples along with measurements of copepod egg production (made using an incubation technique) carried out each week in Long Island Sound during 1985 and a 25-day time-series of daily measurements of the same variables at a fixed station in the nearshore coastal upwelling zone off Concepción, Chile, in January 1986. Egg production was often low, suggesting food-limitation. Temporal variations in the fecundity of the copepods Temora longicornis and Acarlia tonsa in Long Island Sound did not follow changes in total phytoplankton biomass, but rather changes in the concentration of chlorophyll in the > 10 μm and > 20 μm size fractions. Off Chile, changes in fecundity of Calanus chilensis mirrored changes in both total phytoplankton biomass and the > 20 μm size fraction. This fact demonstrates that herbivorous copepods follow closely changes in their food environment and suggests that there is a close coupling between food quality of phytoplankton (in terms of particle size), ingestion and egg production by adult females.     

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.     

Tidal exchange of zooplankton between Lough Hyne and the adjacent coast

Plankton samples collected in November 2002, February, May and August 2003 were used to examine seasonal variation in tidal exchange of zooplankton biomass, abundance and species composition between Lough Hyne Marine Nature Reserve and the adjacent Atlantic coast. Micro- to mesozooplankton were collected by pump over 24-h sampling periods during spring and neap tides from the narrow channel connecting the semi-enclosed water body to the Atlantic. Sample biomass (dry weight) and total zooplankton abundance peaked in the summer and were lowest in winter, showing a positive relationship with temperature. Zooplankton biomass, total abundance and numbers of holo- and meroplankton revealed import during some diel cycles and export in others. However, the tidal import of these planktonic components was generally dominant, especially during May. The greatest import of numbers of holoplankters and meroplanktonic larvae occurred during May and August, respectively. There was no significant variation in sample biomass between periods of light and dark, but some variation in zooplankton abundance could be explained by this diel periodicity. Significant differences in sample assemblage composition between flood and ebb tide samples were always observed, except during winter neap tides. There was a net import of the copepods Temora longicornis and Oithona helgolandica and the larval stages of Mytilus edulis during spring and summer. Proceraea cornuta and Capitellid trochophores were imported during winter, and a hydrozoan of the genus Obelia during the spring spring tides. Seasonal export from the lough was shown by Pseudopolydora pulchra larvae (autumn and spring), Serpulid trochophores (autumn) and veligers of the bivalve Anomia ephippium (summer). It is suggested that the direction of tidal exchange of meroplanktonic taxa is related to the distribution of the adult populations. Copepod naupliar stages dominated the assemblages except during May spring tides when the copepod Pseudocalanus elongatus made up over 22% of the abundance. The general import of micro- to mesozooplankton may, in part, explain the higher densities of this size-class of zooplankton within the semi-enclosed system of Lough Hyne.     

Reproductive success of Calanus pacificus during diatom blooms in Dabob Bay, Washington

While numerous laboratory studies in the last decade have shown that diatoms can induce reproductive failure in copepods, field evidence for a negative diatom effect is equivocal. To unambiguously elucidate the effects diatoms have on copepod reproduction in situ, we undertook a study of the abundance, distribution, grazing rates, and reproductive success of Calanus pacificus in Dabob Bay, Washington, USA, during two spring bloom periods. We simultaneously measured the phytoplankton composition, abundance, and distribution. Here we present results for the reproductive success of C. pacificus using four measurements: egg production rate, clutch size, egg hatching success, and naupliar survival (to the first feeding stage). Egg production rate was positively correlated with chlorophyll a concentration, and egg hatching success and naupliar survival were usually greater than 80%. However, in February 2002 and 2003 – during blooms of diatoms of the genus Thalassiosira – either egg hatching success, naupliar survival, or both were significantly depressed compared to other times in spring and summer. These effects, combined with evidence that C. pacificus was grazing aldehyde-producing Thalassiosira at the time of their blooms, indicate that diatoms can negatively affect copepod reproduction in the field, albeit only under specific circumstances and for brief periods.     

The seasonal cycle of surface chlorophyll in the Peruvian upwelling system: A modelling study

The seasonal variability of surface chlorophyll in the northern Humboldt Current System is studied using satellite data, in situ observations and model simulations. The data show that surface chlorophyll concentration is highest in austral summer and decreases during austral winter, in phase opposition with coastal upwelling intensity. A regional model coupling ocean dynamics and biogeochemical cycles is used to investigate the processes which control this apparently paradoxical seasonal cycle. Model results suggest that the seasonal variability of the mixed layer depth is the main controlling factor of the seasonality. In winter, the mixed layer deepening reduces the surface chlorophyll accumulation because of a dilution effect and light limitation. In summer, biomass concentrates near the surface in the shallow mixed layer and nitrate limitation occurs, resulting in a biomass decrease in the middle of summer. Intense blooms occur during the spring restratification period, when winter light limitation relaxes, and during the fall destratification period, when the surface layer is supplied with new nutrients. Model sensitivity experiments show that the seasonal variations in insolation and surface temperature have little impact on the surface chlorophyll variability.