Copepod grazing during spring blooms: Does Calanus pacificus avoid harmful diatoms?

During late winter and spring of 2002 and 2003, 24, 2–3 day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. The results of the copepod grazing experiments for C. pacificus are discussed here. Each week, copepod grazing incubation experiments from two different depth layers were conducted. Grazing was measured by both changes in chlorophyll concentration and cell counts. In 2002, there was one moderate bloom consisting mainly of Thalassiosira spp. in early February, and a larger bloom in April comprised of two Chaetoceros species and Phaeocystis sp. Similarly, in 2003, there were two blooms, an early one dominated by Thalassiosira spp., and a later one consisting of Chaetoceros spp. and Thalassiosira spp. Clearance rates on individual prey species, as calculated by cell counts, showed that C. pacificus are highly selective in their feeding, and may have much higher clearance rates on individual taxa than rates calculated from bulk chlorophyll disappearance. During weeks of high phytoplankton concentration, the copepods generally ate phytoplankton. However, they often rejected the most abundant phytoplankton species, particularly certain Thalassiosira spp., even though the rejected prey were often of the same genus and similar size to the preferred prey. It is speculated that this avoidance may be related to the possible deleterious effects that certain of these diatom species have on the reproductive success of these copepods. During weeks of medium to low phytoplankton concentration, the copepods selectively ate certain species of phytoplankton, and often had high electivity for microzooplankton. The selection mechanism must consist of active particle rejection most likely based on detection of surface chemical properties, since the diatoms that were selected were of the same genus, nearly the same size, and at lower numerical abundance than those cells that were avoided. The grazing choices made by these copepods may have important consequences for the overall ecosystem function within coastal and estuarine systems through changes in the transfer efficiency of energy to higher trophic levels.     

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.     

Spatio-temporal variability of small copepods (especially Oithona plumifera) in shallow nearshore waters off the south coast of South Africa

Although small copepods are one of the main dietary sources for many commercially important fish, their role in the pelagic trophic dynamics has traditionally been underestimated due to the methodology commonly used in plankton sampling. Temporal variation in abundance of adults and nauplii of small copepods (particularly Oithona plumifera) in nearshore waters on the south coast of South Africa was investigated fortnightly over 14 months at site (km) and location (100 m) scales. Sampling was within <500 m of the shore, where depth was ca. 10 m, using vertical hauls of an 80-μm mesh plankton net from 1 m above the seabed to the surface. Twenty-seven adult copepod taxa were recorded, but Oithona spp. was consistently the most abundant. Taxon richness was 7–19 on each sampling occasion. There was strong temporal variation (Oithona varied between 0 and 2300 m⁻³), but much of this was short-term variability (e.g. between consecutive sampling sessions), with no seasonality or other long-term discernable patterns. There were periods of consistently low numbers, but very high numbers often followed samples with low abundances. Nor was there spatial structure at the location scale, though numbers differed between sites. Despite considerable variability at the location scale within sites, Kenton consistently showed higher densities than High Rocks. Separate analyses, with Bonferroni adjustment, showed that this difference was significant on eight out of 21 occasions for Oithona, six for other pelagic copepods and three for nauplii. This suggests that hydrodynamics favour aggregation of plankton at Kenton. A high degree of short-term variability, with a tendency for aggregation of small zooplankton at certain sites has implications for both pelagic processes and food-web links between the benthic and pelagic environments.     

Plankton response to El Niño 1997–1998 and La Niña 1999 in the southern region of the California Current

The IMECOCAL Program began in 1997, with the objective of sampling plankton systematically in the Mexican region of the California Current. We present results of chlorophyll a concentrations and zooplankton displacement volumes for the eight cruises from September 1997 to October 1999. The abundance of 22 zooplankton groups was also analyzed for the first four cruises. The response of plankton to the 1997–1998 El Niño was atypical. From September 1997 to January 1998, chlorophyll a and zooplankton volume were at typical values (median integrated chlorophyll was 27 mg/m² and zooplankton 100 ml/1000 m³ in 9801/02). After the peak of El Niño, the system shifted to cooler conditions. Integrated chlorophyll gradually increased to a median of 77 mg/m² in April 1999. In contrast, zooplankton volumes decreased from October 1998 onward, despite favorable phytoplankton availability in 1999. Zooplankton structure was dominated by copepods and chaetognaths through the ENSO cycle, but interannual changes were evident. In the fall of 1997 there was a higher proportion of copepods, chaetognaths, and other minor groups, while the fall of 1998 zooplankton was richer in salps and ostracods. Historical data from previous Baja California CalCOFI cruises indicated that zooplankton volumes measured during the IMECOCAL cruises were above the long-term mean for the period 1951–1984. This suggests a differential response of plankton to the El Niño of 1997–1998 compared to the El Niño of 1957–1959. Regional differences in zooplankton volumes were also found, with central Baja California having 41% higher biomass than northern Baja California. Volumes from both regions were larger than those recorded by CalCOFI off southern California during 1997–1998, but the situation was reversed in 1999. The higher biomasses in the 1997–1998 El Niño can be attributed to high abundance of salps, which showed an affinity with warm, saline water.     

Copepod grazing during spring blooms: Can Pseudocalanus newmani induce trophic cascades?

During late winter and spring of 2002 and 2003, 24 two- to three-day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. Here, we discuss the results of our grazing experiments for P. newmani. Each week, we conducted traditional microzooplankton dilution experiments and “copepod dilution” experiments, each from two different layers. Grazing was measured by changes in chlorophyll concentration and direct cell counts. Clearance rates on individual prey species, as calculated by cell counts, showed that Pseudocalanus are highly selective in their feeding, and may have much higher grazing rates on individual taxa than calculated from bulk chlorophyll disappearance. The grazing rates of the copepods, however, are typically an order of magnitude lower than the grazing rates of the microzooplankton community, or the growth rates of the phytoplankton. P. newmani ingested diatoms, but, at certain times fed preferentially on microzooplankton, such as ciliates, tintinnids, and larger dinoflagellates. Removal of the microzooplankton may have released the other phytoplankton species from grazing pressure, allowing those species’ abundance to increase, which was measured as an apparent “negative” grazing on those phytoplankton species. The net result of grazing on some phytoplankton species, while simultaneously releasing others from grazing pressure resulted in bulk chlorophyll-derived estimates of grazing which were essentially zero or slightly negative; thus bulk chlorophyll disappearance is a poor indicator of copepod grazing. Whether copepods can significantly release phytoplankton from the grazing pressure by microzooplankton in situ, thus causing a trophic cascade, remains to be verified, but is suggested by our study.     

Grazing impact of mesozooplankton in an upwelling region off northern California, 2000–2003

Mesozooplankton (>200 μm) grazing impact (% phytoplankton standing crop consumed d⁻¹) was determined by the gut fluorescence method during three springs (2000, 2001 and 2002) and two winters (2002 and 2003) in a coastal upwelling region off northern California. Wind events, in terms of both magnitude and duration, varied inter-annually and seasonally and included both upwelling-favorable and relaxation events. Grazing impact of mesozooplankton also varied inter-annually and seasonally, and was highest during June 2000 (mean=129% of standing crop d⁻¹), a prolonged period of wind “relaxation” and phytoplankton bloom. In contrast, mean grazing impact was lower during periods of stronger, more persistent winds, more active upwelling, greater cross-shelf transport, and lower chlorophyll concentration (25% and 38% in May–June 2001 and 2002, respectively). Wintertime conditions (January 2002 and 2003) were characterized by weakly upwelling or downwelling-favorable winds, low chlorophyll concentration, and lower mean mesozooplankton grazing impact (13% and 12%, respectively). The larger (>500 μm) size class contributed proportionally more to total mesozooplankton (>200 μm) grazing impact than the smaller (200–500 μm) size class during all sampling periods except spring 2002. These results suggest that mesozooplankton grazing impact is higher in spring than in winter, and that during the spring upwelling season, grazing is higher during periods of wind relaxation (weak upwelling) than during periods of stronger upwelling. Further, these results suggest an important role of mesozooplankton grazers on phytoplankton dynamics in the upwelling region off northern California.     

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.     

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.     

Feeding Strategy of Japanese Sand Lance Larvae in Relation to Ciliated Protozoa in the Vicinity of a Thermohaline Front

Larval feeding and survival strategies are described on a Japanese sand lance, Ammodytes personatus Girard, collected in the vicinity of a thermohaline front in the Ise Bay, in comparison with those of other predominant larvae, Hexagrammos spp. and Sebastiscus marmoratus (Cuvier). First-feeding A. personatus larvae (3.1–3.9 mm NL) fed primarily on tintinnid ciliates, subsequently switching to copepod nauplii (4.0–7.9 mm NL larvae) and post-naupliar copepods (8.0–11.3 mm NL larvae). First-feeding Hexagrammos spp. larvae (6.5–6.9 mm NL) fed primarily on post-naupliar copepods, and first-feeding S. marmoratus larvae (<4.0 mm NL), mostly on copepod nauplii. The different food preferences of these species at first-feeding were related to their different mouth widths (0.15–0.19, 0.52–0.56 and 0.32–0.40 mm, respectively) and/or body size (3.1–3.9, 6.5–6.9 and 3.2–3.9 mm NL, respectively). Ciliate-feeding by first-feeding A. personatus larvae was strongly related to the convergence of the larvae and their prey near the thermohaline front, densities of both being greater on the inshore side of the frontal zone. In conclusion, the aggregation of ciliates near the thermohaline front may have improved feeding conditions and survival of first-feeding A. personatus larvae.     

Association of picophytoplankton distribution with ENSO events in the equatorial Pacific between 145°E and 160°W

Climatological variability of picophytoplankton populations that consisted of >64% of total chlorophyll a concentrations was investigated in the equatorial Pacific. Flow cytometric analysis was conducted along the equator between 145°E and 160°W during three cruises in November–December 1999, January 2001, and January–February 2002. Those cruises were covering the La Niña (1999, 2001) and the pre-El Niño (2002) periods. According to the sea surface temperature (SST) and nitrate concentrations in the surface water, three regions were distinguished spatially, viz., the warm-water region with >28 °C SST and nitrate depletion (<0.1 μmol kg⁻¹), the upwelling region with <28 °C SST and high nitrate (>4 μmol kg⁻¹) water, and the in-between frontal zone with low nitrate (0.1–4 μmol kg⁻¹). Picophytoplankton identified as the groups of Prochlorococcus, Synechococcus and picoeukaryotes showed a distinct spatial heterogeneity in abundance corresponding to the watermass distribution. Prochlorococcus was most abundant in the warm-water region, especially in the nitrate-depleted water with >150×10³ cells ml⁻¹, Synechococcus in the frontal zone with >15×10³ cells ml⁻¹, and picoeukaryotes in the upwelling region with >8×10³ cells ml⁻¹. The warm-water region extended eastward with eastward shift of the frontal zone and the upwelling region during the pre-El Niño period. On the contrary, these regions distributed westward during the La Niña period. These climatological fluctuations of the watermass significantly influenced the distribution of picophytoplankton populations. The most abundant area of Prochlorococcus and Synechococcus extended eastward and picoeukaryotes developed westward during the pre-El Niño period. The spatial heterogeneity of each picophytoplankton group is discussed here in association with spatial variations in nitrate supply, ambient ammonium concentration, and light field.     

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.     

Feeding of copepods on natural suspended particles in Tokyo Bay

Community grazing rates of copepods were estimated from data taken during three cruises in Tokyo Bay, based on bottle incubations and a temporal variation of gut fluorescence. Special attention was paid to the feeding selectivity in the estimations. Differential grazing was observed in the copepod communities:Acartia omorii, abundant in February, selectively fed on the particles of dominant size classes, whileOithona davisae, dominant throughout the year, andCentropages abdominalis selected large particles (>20µm). The maximum filtering rates on certain size classes were several times the average. In addition, a 34-hr investigation of the gut fluorescence of copepods revealed nocturnal feeding inParacalanus spp.,Pseudodiaptomus marinus andOithona davisae.Copepod communities collected with a net (95-µm mesh opening) were estimated to graze, in February 3.0%, in August 3.1–4.5% and in November 4.2–11.9% of the standing crops of phytoplankton or suspended particles per day.     

Resource Partitioning Among Planktivorous Fish Larvae and Fry in a Mediterranean Coastal Lagoon

Resource partioning among the planktivorous stages of eight fish species occurring in a Mediterranean coastal lagoon was studied. Five species were migratory mullets that spawned in the sea (Liza ramada, L. aurata, L. saliens, Chelon labrosusandMugil cephalus), while the other three species were resident and spawned in the lagoon (Atherina boyeri, Cyprinus carpioandGambusia holbrooki).Mullet fry exhibited similar diets, based on the consumption of zooplanktonic Crustacea such as copepods and cladocerans, although adult chironomids were also important.Gambusia holbrookiandA. boyerifed on small prey such as copepod nauplii, copepodites and rotifers, whileC. carpioconsumed larger prey preferentially (cladocerans and copepods).According to the trophic overlap and niche width results, seven species pairs could be competing in the estuary, although further experimental evidence is still required.     

High-West and Low-East Distribution Patterns of Chlorophyll a, Primary Productivity and Diatoms in the Subarctic North Pacific Surface Waters, Midwinter 1996

We have determined chlorophyll a (Chla) concentration, primary productivity, cell density and species composition of diatoms, and the number of microzooplankton at the surface in the subarctic North Pacific in January 1996. The wet weight of copepods obtained by vertical tows from 150 m to the surface was also measured during the cruise. Chla concentration and primary productivity tended to be higher in the region west of 180°, the western subarctic North Pacific (WSNP), than east of 180°, the eastern subarctic North Pacific (ESNP). The same results were observed for the total diatom cell densities and for the genera Thalassiosira and Denticulopsis. Significant linear relationships were observed between the Thalassiosira cell density and Chla concentration and primary productivity, indicating that Thalassiosira contributes to the high-WSNP and low-ESNP distribution patterns of Chla concentration and primary productivity. Moreover, naked ciliate abundance tended to be lower in the WSNP than in the ESNP, whereas copepod biomass showed an inverse trend. Significantly negative Spearman rank correlations were found between the Thalassiosira cell density and the number of naked ciliates and between the number of naked ciliates and the wet weight of copepods. These results indicate that copepod grazing indirectly controls Thalassiosira cell density via predation on the naked ciliates. We conclude that the high copepod biomass in the WSNP is a factor controlling the high-WSNP and low-ESNP Thalassiosira abundance and hence Chla concentration and primary productivity patterns.     

Occurrence of copepod carcasses in the lower Chesapeake Bay and their decomposition by ambient microbes

We tested and refined the Neutral Red staining method for separating live and dead copepods in natural samples. Live copepods were stained red whereas dead copepods remained unstained. The staining results were not affected by method of killing, time of death or staining time. Tow duration had no significant effect on the percent dead copepods collected. The Neutral Red staining method was applied to study the occurrence of dead copepods along the York River and the Hampton River in the lower Chesapeake Bay during June–July, 2005. The zooplankton community was dominated by copepods; on average 29% of the copepod population appeared dead. Recovery of percent dead copepods did not differ between horizontal tows and vertical tows, suggesting that dead copepods were homogenously distributed in the water column. No significant relationship was found between the percent dead copepods and surface water temperature, salinity, Secchi depth or chlorophyll concentration. In laboratory experiments, dead copepods were decomposed by ambient bacteria and the rate of decomposition was temperature-dependent. Combining field and laboratory results we estimated that the non-consumptive mortality (mortality not due to predation) of copepods in the lower Chesapeake Bay was 0.12 d⁻¹ under steady-state condition, which is within the global average of copepod mortality rate.     

Coupling of life cycles of the copepods Calanus chilensis and Centropages brachiatus to upwelling induced variability in the central-southern region of Chile

A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from August 2002 to December 2003 allowed the study of annual life cycles of the copepods Calanus chilensis and Centropages brachiatus in association with environmental variability in the coastal upwelling zone. Changes in the abundance of eggs, nauplii, and copepodids were assessed from samples taken at a mean time interval of ca. 20 days. Upwelling variability in near-surface waters was reflected in seasonal changes in salinity, water column stratification, and oxycline depth, as well as a weak seasonal signal in sea surface temperature (1-2 °C). Both copepods exhibited similar life cycles, characterized by continuous reproduction throughout the year. Estimates of generation times, as a function of temperature, were 25-30 days for C. chilensis and 27-35 days for C. brachiatus, predicting about 12 and 10 generations a year, respectively. These estimates were consistent with reproduction pulses observed in the field. It was thus suggested that copepods may grow under non-limiting food conditions in this upwelling area. However, despite continuous reproduction, there were abrupt changes in population sizes along with the disappearance of early naupliar and copepodid stages taking place even during the upwelling season (spring/summer). These changes were attributed to sudden increases in mortality taking place in spring or early summer, after which the populations remained at low levels through the fall and winter. It is thus suggested that, in addition to variability in the physical environment, biological interactions modulating changes in copepod mortality should be considered for understanding copepod life cycles in highly productive upwelling systems.     

The contribution of microzooplankton to the diet of mesozooplankton in an upwelling filament off the north west coast of Spain

Incubation experiments were carried out daily during a Lagrangian experiment within an upwelled filament off the Galician coast to determine the importance of microzooplankton in the diet of calanoid copepods. Despite low chlorophyll concentrations the microzooplankton formed the minor component of the diet of the copepod community (7 to 15% of carbon ingested through autotrophic and heterotrophic prey). Ingestion of ciliates was greater than that of heterotrophic dinoflagellates, which reflected a higher abundance of ciliates in the water column. Heterotrophic nanoflagellates appeared also to be consumed by the copepods, although the very small size fraction (2–5μm) was probably not grazed by the larger copepods of Calanus spp. Grazing pressure by the copepods enumerated in the net samples was not sufficient to impact significantly the microzooplankton populations (2 to 51% of daily microzooplankton production was removed). Allometric relationships of grazing on microzooplankton for a range of numerically dominant copepod species are developed from the experimental results. The grazing pressure of the whole copepod community is estimated from these relationships. By considering the total mesozooplankton community we suggest that microzooplankton growth was probably restricted by metazoan grazers.     

Production of <Emphasis Type="Italic">Acartia steueri</Emphasis> (Copepoda: Calanoida) in Ilkwang Bay,Southeastern Coast of Korea

Production of the marine calanoid copepod Acartia steueri was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay, on the southeastern coast of Korea. Phytoplankton standing stock ranged over 1.0 to 9.3 mg chl.a m⁻³, and annual primary productivity (by the C-14 method) at three stations was estimated at 200 gC m⁻² yr⁻¹. Acartia steueri (nauplii + copepodids + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. steueri, excluding the NI stage, was 0.01–4.55 mgC m⁻³ (mean: 0.68 mgC m⁻³) with peaks in November, February, May and July-early August, and relatively low biomass in September– January. Instantaneous growth rates of the nauplius stages were higher than the copepodid stages. Annual production of A. steueri was 25.1 mgC m⁻³ yr⁻¹ (or 166 mgC m⁻² yr⁻¹), showing peaks in November, May and July–August with a small peak in February, and low production in December–April and September–October. There were no significant relationships between the daily production rate of A. steueri and temperature or chlorophyll a concentration, indicating that unknown other factors might be related to the variation of the production rate.     

Surface distributions of copepods in relation to regional upwellings around the Izu Islands in summer of 1988

Surface distributions of zooplankton were surveyed with simultaneous measurements of temperature, salinity, chlorophyll and nitrate, around the Izu Islands in July 1988. Several phytoplankton patches, which had developed in the upwelled waters, were encountered around the islands. Zooplankters collected with a 20µm mesh mainly consisted of copepods including eggs, nauplii, copepodites and adults.Paracalanus parvus dominated among the copepod females. High concentrations of all copepod stages were associated with the phytoplankton patches. The increase in reproductive activity of female copepods was considered as a possible process to form the copepod patches associated with the phytoplankton patches.     

Seasonal and spatial variation in abundance and egg production of Paracalanus parvus (Copepoda: Calanoida) in/out Jiaozhou Bay, China

The spatial distribution of stage-specific abundance and reproduction of the copepod Paracalanus parvus were studied from October 2005 to September 2006 in the Jiaozhou Bay. This copepod occurred continuously in this bay throughout the year. The species reached the lowest abundance in April and peaked in June. From October to December, distribution center mainly occurred in offshore water and at the mouth of the bay. In winter, early copepodites and adults gradually decreased and till February, most of the population was only comprised of CIV–CV stages. Overwintering copepodites matured in March and males tended to mature before female. From May to September, each stage occurred in the population and gradually reached high abundance. Temperature and chlorophyll a (Chl-a) concentration in the three stations can't clearly explain the seasonal variation in stage-specific abundance, so we surmised the important effect of the Yellow Sea. Egg production rate (EPR) reached its lowest in winter and peaked in June at 60.8 eggs female⁻¹ day⁻¹ in nearshore water. In the warming period, EPR in nearshore water was statistically higher and EPR > 10 eggs female⁻¹ day⁻¹ lasted longer than that in offshore water, showing the importance of nearshore water for recruitment of P. parvus. Our study showed that EPR was positively related to temperature and total chlorophyll a in offshore water and mouth of the bay. In nearshore water, the relationships between EPR and temperature and Chl-a in three size fractions were not the same as those in offshore water, suggesting complicated ecosystem in such a eutrophic area in warming period.