Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

The predation impact of juvenile herring Clupea harengusand sprat Sprattus sprattus on estuarine zooplankton

The consumption of estuarine copepods by juvenile herring and sprat during estuarine residency was estimated using fish biomass data and daily rations calculated from two models of feeding in fish: a bioenergetic model and a gastric evacuation model. The bioenergetic model predicted daily rations that were, on average, three times higher than those estimated by a model based on field records of stomach contents. The biomass of herring and sprat in the estuary was negatively correlated with the daily ration suggesting that the clupeid fish populations were resource-limited. Copepod production decreased towards the winter and peaked in spring and summer. The relative importance of predation changed seasonally in function of the migration pattern of herring and sprat. In the spring and the summer, in situ production ofcopepod biomass was higher than the in situ consumption by fish. During the fall and the winter, consumption exceeded production. This suggests that top–down control exerted by marine pelagic fish may be an important force structuring estuarine copepod populations.     

Composition,distribution and biomass of meiobenthos in the Oosterschelde estuary (SW Netherlands)

Meiofauna composition, abundance, biomass, distribution and diversity were investigated for 31 stations in summer. The sampling covered the whole Oosterschelde and comparisons between the subtidal — intertidal and between the western-central — eastern compartment were made.Meiofauna had a community density ranging between 200 and 17 500 ind 10 cm^–2, corresponding to a dry weight of 0.2 and 8.4 gm^–2. Abundance ranged between 130 and 17 200 ind 10 cm^–2 for nematodes and between 10 and 1600 ind 10 cm^–2 for copepods. Dry weight biomass of these taxa was between 0.5–7.0 gm^–2 and 0.008–0.3 gm^–2 for nematodes and copepods respectively.The meiofauna was strongly dominated by the nematodes (36–99%), who's abundance, biomass and diversity were significantly higher intertidally than subtidally and significantly higher in the eastern part than in the western part. High numbers were positively correlated with the percentage silt and negatively with the median grain size of the sand fraction. The abundance and diversity of the copepods were highest in the subtidal, but their biomass showed an inverse trend being highest on the tidal flats.The taxa diversity of the meiofauna community and species diversity of both the nematodes and the copepods were higher in subtidal stations than on tidal flats. In the subtidal, the meiofauna and copepod diversity decreased from west to east, whereas nematode diversity increased.The vertical profile clearly reflected the sediment characteristics and could be explained by local hydrodynamic conditions.Seasonal variation was pronounced for the different taxa with peak abundance in spring, summer or autumn and minimum abundance in winter.Changes in tidal amplitude and current velocity enhanced by the storm-surge barrier will alter the meiofauna community structure. As a result meiofauna will become more important in terms of density and biomass, mainly due to increasing numbers of nematodes, increasing bioturbation, nutrient mineralisation and sustaining bacterial growth. A general decrease in meiofauna diversity is predicted. The number of copepods is expected to decrease and interstitial species will be replaced by epibenthic species, the latter being more important in terms of biomass and as food for the epibenthic macrofauna and fishes.     

Long-term changes in the population of Eurytemora affinis (Copepoda,Calanoida) in the Gironde estuary (1978–1992)

Long-term changes in the population of Eurytemora affinis were investigated during 15 years (1978–1992) at a fixed station in the Gironde estuary (South West France). Total numbers, sex-ratio, % ovigerous females, proportion of copepodites and clutch-size were taken into account and their long-term distribution was related to temperature, river flow, salinity, suspended matter and chlorophyll a concentrations after the seasonal effect was removed for all variables.There was a great decrease of the river flow during the period of investigation, due to a general deficit in pluviosity. On the contrary, the general trend for salinity was an increase from 1978 to 1992. The same pattern was observed for chlorophyll, indicating an intrusion of marine phytoplankton. Water temperature increased significantly during the study period, following the general increase in air temperature observed in the area. For the suspended particulate matter (SPM), an increase was observed between 1978 and 1981–1982, then a very sharp decrease occurred from 1984 onwards.Numbers of E. affinis were inversely correlated with temperature and salinity, and positively correlated with the river flow. E. affinis was negatively correlated with chlorophyll concentration because of the covariation with salinity. No clear long-term trend was observed for the sex-ratio. A significant correlation was found between females carrying egg-sacs and SPM concentration, probably due to a decrease of the predation pressure in very turbid waters. The percentages of copepodites tended to decrease with time and were inversely correlated with temperature. Clutch-size significantly decreased during the 15-year period. This trend was mainly explained by temperature and salinity.It is concluded that, in the absence of strong human alteration in the Gironde estuary the long-term distribution of zooplankton can be explained by the natural environmental variability. The dominant factors are the river flow which governs the movement of the populations along the estuary and the temperature which influences the reproductive processes. Any change in climate will have consequences on the pelagic community.     

Space and time distribution of zooplankton in a Mediterranean lagoon (Etang de Berre)

In the Berre lagoon, a large brackish and swallow area near Marseille, the environmental factors (temperature, salinity, oxygen, suspended particulate matter and chlorophyll) generally display strong space and time variations. The rotifer Brachionus plicatilis and the copepod Acartia tonsa constitute the bulk of the zooplankton population during all the year. Their space and time distributions were studied in 23 stations distributed all over the lagoon, during four seasonal cruises (February, June, October, November), at surface and bottom layers. There is no marked difference in the horizontal and vertical distribution of the two species, (except in November when rotifers were prevailing in surface and copepods at depth) and in their time occurence. When the four series of data are pooled, correlation analysis show that A.tonsa is positively correlated with temperature, salinity and seston and negatively to oxygen and chlorophyll. B. plicatilis is positively correlated with temperature and seston, but also with chlorophyll, while salinity has a negative effect. The specific eggs number of both species is chlorophyll dependent. Considering seasonal cruises separately, some differences appear in the sense or the significance of these different correlations. The respective distribution of the two species is only partly dependent on the variation of the environmental factors: most of the variance remains unexplained, as indicated by the result of a stepwise multiple regression analysis using the most significant factors (temperature, salinity and oxygen explain 33 to 42% of the variance in Acartia, while temperature and salinity explain 27 to 28% of the variance in Brachionus). Thus, internal behavioral factors could also play a role in the distribution of organisms, particularly in some cases of aggregations of organisms observed during this study. As the two species occupied the same space habitat most of the year, they are potentially in competition for food. A way to optimize the food utilization could be the time separation of their feeding activity, nocturnal in Acartia and diurnal in Brachionus. Another way could be selective feeding upon food particles depending on their size (Brachionus being able to use finer particles than Acartia) or their quality (Brachionus being more herbivorous than Acartia) as demonstrated in some grazing experiments carried out in parallel.     

Feeding rates and productivity of the copepod Acartia bifilosa in a highly turbid estuary; the Gironde (SW France)

Acartia spp. are the dominant copepod species in the Gironde estuary, seaward of the turbidity maximum area. Acartia bifilosa develop a large population in spring and early summer whereas Acartia tonsa appear in late summer. High values and high variability of chlorophyll a/suspended particulate matter ratio are found seaward of the turbidity maximum area. Feeding rates of A. bifilosa were measured by fluorometry. Phytoplankton ingestion was found to be highly variable, between 8 to 80% of copepod carbon body weight. Except for adult females, copepods were heavier in summer than in winter. PB ratios, estimated by the instantaneous growth rate method, varied from 0.03 d^–1 to 0.14 d^–1. The gut contents and P/B ratios of Acartia bifilosa were related to chl a/SPM ratio. From those data, and a few obtained for A. tonsa, it is concluded that only in summer months phytoplankton ingestion is enough to maintain secondary production.     

A marine calanoid copepod (Temora longicornis) in a semi-stagnant coastal lagoon (Lake Grevelingen,S.W.-Netherlands)

The marine calanoid copepodTemora longicornis was introduced in saline Lake Grevelingen (S.W.-Netherlands), when the lake was continuously flushed with North Sea water in 1979, to enhance the reduced salinity. Development of the copepod proceeded suboptimally as a consequence of an unfavourable environment in the lake: 1. The phytoplankton assemblage, mainly consisting of crytomonad flagellates, less suitable for growth and reproduction of the adult stages. 2. An acute food shortage occurring at a crucial moment in the development. 3. A. pronounced salinity-and oxygen stratification in June, preventing further development ofTemora.Communication nr. 474 of the Delta Institute for Hydrobiological Research, Yerseke.     

Zooplankton species composition in the Oosterschelde (SW Netherlands) before,during and after the construction of a storm-surge barrier

During the period of construction of a storm-surge barrier current velocities decreased strongly and the Eastern compartment of the basin obtained a lagoon-like character. The rotifer Synchaeta spp., already abundant in the neighbouring salt and brackish lakes, profited from this condition. Higher and less fluctuating salinities caused the estuarine character of this compartment to disappear in the post-barrier years. This was reflected in the obscured succession of some Acartia species: the estuarine A. tonsa dominated in pre-barrier- and barrier years during summer, while the marine A. clausi was abundant in the post-barrier Oosterschelde during that time. Also the changed food conditions (Bakker & Vink, 1993) may have played a role in this phenomenon.Further changes in species composition were hardly observed. The main changes were of a quantitative nature (Bakker & Van Rijswijk, 1993; Tackx et al., 1993). A list of the commonly occurring species is given.     

Fitness consequences for copepods feeding on a red tide dinoflagellate: deciphering the effects of nutritional value, toxicity, and feeding behavior

Phytoplankton exhibit a diversity of morphologies, nutritional values, and potential chemical defenses that could affect the feeding and fitness of zooplankton consumers. However, how phytoplankton traits shape plant–herbivore interactions in the marine plankton is not as well understood as for terrestrial or marine macrophytes and their grazers. The occurrence of blooms of marine dinoflagellates such as Karenia brevis suggests that, for uncertain reasons, grazers are unable to capitalize on, or control, this phytoplankton growth—making these systems appealing for testing mechanisms of grazing deterrence. Using the sympatric copepod Acartia tonsa, we conducted a mixed diet feeding experiment to test whether K. brevis is beneficial, toxic, nutritionally inadequate, or behaviorally rejected as food relative to the palatable and nutritionally adequate phytoplankter Rhodomonas lens. On diets rich in K. brevis, copepods experienced decreased survivorship and decreased egg production per female, but the percentage of eggs that hatched was unaffected. Although copepods showed a 6–17% preference for R. lens over K. brevis on some mixed diets, overall high ingestion rates eliminated the possibility that reduced copepod fitness was caused by copepods avoiding K. brevis, leaving nutritional inadequacy and toxicity as remaining hypotheses. Because egg production was dependent on the amount of R. lens consumed regardless of the amount of K. brevis eaten, there was no evidence that fitness costs were caused by K. brevis toxicity. Copepods limited to K. brevis ate 480% as much as those fed only R. lens, suggesting that copepods attempted to compensate for low food quality with increased quantity ingested. Our results indicate that K. brevis is a poor food for A. tonsa, probably due to nutritional inadequacy rather than toxicity, which could affect bloom dynamics in the Gulf of Mexico where these species co-occur.     

Is the copepod egg production in a highly turbid estuary (the Gironde, France) a function of the biochemical composition of seston?

The influence of the biochemical composition of particles originating from surface waters of the Gironde estuary on egg production rates of Eurytemora affinis zooplanktonic population was studied. In the high turbidity zone, suspended particulate matter had a low nutritional quality because the easily available organic fraction represented less than 15% of the overall particulate organic matter. In waters located seaward of the high turbidity zone, a slight increase in nutritional quality was observed. As a result, the sum of easily extractable organic macromolecules represented 15 to 33% of the overall particulate organic matter. The present study suggests that the low egg-production rate of Eurytemora affinis, occurring in the high turbidity zone, results from combined effects of temperature and bad feeding conditions in the area. Low copepod production can be explained by little phytoplankton growth due to light limitation and, therefore, restricted food availability, as well as difficulties in food selection, non-living particle may being dominant.     

Feeding behaviour and migrations in a natural population of the copepod Acartia tonsa

This paper deals with the variations on feeding activities and diel migrations of Acartia tonsa Dana, the dominant copepod species in Berre lagoon (west Mediterranean French coasts).A 27 hour in situ study was carried out during June 1989, at a station located in the south west of the lagoon. Vertical profiles of salinity, temperature and dissolved oxygen taken each 12 hours showed a stratification of the water column in two distinct layers: (1) a superficial layer with higher temperature, moderate salinity, and high oxygen concentrations; (2) a colder, more saline and almost anoxic deep layer. In situ chlorophyll a measurements were made at –1, –3 and –5 m; concentrations were relatively homogeneous through the water column during the whole sampling period.Zooplankton samples were collected every 3 hours with a 200 µm mesh net, in three strata (0–2 m, 2–4 m, 4–6 m). A complete dominance of A. tonsa was observed in the zooplankton community. Our results point out clearly a nocturnal migration, with individuals concentrating in both superficial layers; thus an unimodal migratory pattern can be inferred. Gut flourescence measured following the Mackas & Bohrer technique (1976), showed higher values during night time, and values for females were the highest with wider day—night variations. Similar results were found in laboratory experiments with copepods fed with a culture of Dunaliella tertiolecta.Gut evacuation rate was measured in two laboratory experiments either mixing or separating males and females. Evacuation rate was 17.92 and 27.25 min for males and females respectively.Phytoplankton daily ration for A. tonsa calculated by gut flourescence and gut evacuation rate was particularly low, for it represents only 10% of the individual carbon weight.Moreover, grazing impact on phytoplankton is very restricted, it represents less than 1% of the daily phytoplankton stock.     

Production and ecology of eelgrass (Zostera marinal L.) in the Grevelingen estuary,the Netherlands,before and after the closure

The Grevelingen estuary was cut off from the North Sea and from the influences of the river Rhine by a dam in 1971, and became a stagnant salt-water lake. Production and ecology ofZostera marina L. were studied in 1968 and in 1973–1975, both through standing stock estimations, biomass increases in permanent quadrats, and correlation of distribution patterns with ecological factors. After the closure of the estuary the intertidal eelgrass population extended downwards to 5 m below lake level, probably owing to the increased transparency of the water; the area occupied, and the density of the eelgrass beds increased strongly. Eelgrass annual overground production, based on doubled maximum standing crop values in July–August, was estimated at 50 g C/m² in 1968, 121 g C/m² in 1973 and 91 g C/m² in 1975 inZostera beds, and 4 g C/m² in 1968, 18 g C/m² in 1973 and 23 g C/m² in 1975 for the entire Grevelingen area. A minimum estimate of net production inZostera beds at a depth of 0.50–0.75 m, based on short term changes in biomass in 2 permanent quadrats in 1974 and 1975, was 40.5 g C/m²/yr for overground parts and 12.7 g C/m²/yr for underground parts. Horizontal distribution of celgrass is not primarily limited by grainsize distribution, but more by exposure to wave action and currents. On account of irradiance reduction light is a limiting factor in the vertical distribution of the eelgrass population in Lake Grevelingen. Communication no. 146 of the Delta Institute for Hydrobiological Research, Yerseke, The Netherlands.     

The Role of the Predaceous Copepod Parabroteas Sarsi in the Pelagic Food Web of a Large Deep Andean Lake

Parabroteas sarsi is a predaceous calanoid copepod that inhabits both shallow temporary fishless ponds and deep fish lakes of Patagonia and Antarctica. The aim of this study was to analyse the effect of P. sarsi on the plankton structure of a deep Andean lake (>100 m depth) and the zooplankton vertical distribution in order to asses a possible vertical refuge of the predatory copepod against visual fish predation. We tested the extent to which the trophic cascade effect of this predator propagates through the food web. We carried out a vertical sampling in Lake Rivadavia (Patagonia, Argentina) in order to assess zooplankton distribution. P. sarsi showed a vertical distribution towards deeper layers of the water column both at midday and at night, indicating that the copepod had an effective refuge against visual predation. Additionally, we carried out both field and laboratory experiments with the presence of P. sarsi. The predator was observed to affect significantly the survival of the copepod Boeckella michaelseni both in laboratory and field experiments. On the contrary, rotifers and adults of Daphnia cf. commutata were not substantially affected by the predator. B. michaelseni mouthparts revealed an omnivorous diet; therefore a broad phytoplanktonic size spectrum could be affected by this copepod. However, no cascade effect was observed due to the presence of P. sarsi despite the decrease of B. michaelseni abundance.     

Phytoplankton ecology in an unusually stable environment (Montezuma Well,Arizona, U.S.A.)

Relatively minor annual amplitudes of change in certain major nutrients, and especially pH and water temperature were measured in the spring-fed system of Montezuma Well, Arizona during a four year study. phytoplankton diversity was low but for the most part, composition was spatially and temporally constant; total seasonal phytoplankton density was significantly correlated with regional incident light. Phytoplankton species composition changed briefly during and for a short period following the summer monsoon. Ultraplankton (<5 µm diam.) numerically comprised nearly 80% of the phytoplankton community throughout most of the year. The limited residence time of water in the Well may have provided a competitive advantage for cells with high surface area:volume ratios and correspondingly rapid division rates. Nannochloris bacillaris Naum. and Coccomyxa minor Skuja were perennial dominants. Diatom populations did not increase with annual increases in vernal solar radiation. Low pH, high dissolved CO₂, and limited residence time for metabolic inhibitors are considered to be largely responsible for the reduced blue-green populations in the Well. The only flagellated photosynthetic group present in Montezuma Well was the Cryptophyta. Desmid populations were minimal, even though pH was consistently below circumneutral (6.5) and free CO₂ concentrations high. The role of grazing by an amphipod, Hyalella montezuma, on annual phytoplankton abundance is examined.     

Zooplankton biomass in the Oosterschelde (SW Netherlands) before,during and after the construction of a storm-surge barrier

The hydrodynamic consequences of large coastal engineering (barrier-construction) works in the Oosterschelde were: prolonged residence times of the water, increased sinking of particulate material, and higher water transparencies. This strongly influenced the phytoplankton (Bakker et al., 1990; 1994) and phytoplankton biomass increased in the shallow Eastern compartment of the Oosterschelde (Bakker & Vink, 1994) while phytoplankton concentration of the seston rose.Zooplankton biomass, especially of copepods (Temora) and meroplankton (barnacle larvae) increased during the post-barrier period in the eastern compartment. It is hypothesized that this is caused by the improved feeding conditions and the increased retention times in this area.The barrier years 1985 and 1986 were characterized by low current velocities. In the Eastern compartment, this may have favoured the development of the rotifer Synchaeta (Bakker, 1994) and of the important copepod predator Pleurobrachia (Ctenophora).In the Western compartment, zooplankton developments in the post-barrier years were rather similar to those in the pre-barrier period. This led to the disappearance of the previously existing biomass gradients West-East (maxima in West). At present a trend in the opposite direction (maxima in East) is observed.     

Twentyfive years of changes in the distribution and biomass of eelgrass,Zostera marina,in Grevelingen lagoon,The Netherlands

The wax and wane of the eelgrass (Zostera marina L.) population in Grevelingen lagoon (East Atlantic; The Netherlands) has been documented for over 25 years, together with quantitative and semi-quantitative data on environmental variables. The population expanded after the closure of the Grevelingen estuary in 1971, but declined from 4600 ha surface area in 1978 to less than 100 ha in 1993. There is little causal evidence which factors are responsible for the observed dynamics of the population. The incomplete picture emerging from the data is that of an extremely impoverished eelgrass population, living under constant oligo-mesotrophic marine conditions. Both the sexual and the vegetative modes of reproduction are severely stressed by environmental variables, most likely a combination of low temperatures, high salinity, low dissolved silicate and low ammonium concentrations. Survival of the population asks for the restoration of moderate estuarine conditions.Contribution No. 2180 of the Netherlands Institute of Ecology, Nieuwersluis, The Netherlands.     

Effects of large Saduria entomon (Isopoda) on spatial distribution of their small S. entomon and Monoporeia affinis (Amphipoda) prey

We performed laboratory experiments to investigate the effects of predator avoidance and numerical effects of predation on spatial distribution of small Saduria entomon (Isopoda) and Monoporeia affinis (Amphipoda), with large S. entomon as predators. The horizontal distribution and mortality of the prey species, separately and together, were studied in aquaria with a spatial horizontal refuge. We also estimated effects of refuge on mortality of small S. entomon and M. affinis by experiments without the refuge net. In addition, we investigated whether predation risk from large S. entomon influenced the swimming activity of M. affinis, to clarify the mechanisms behind the spatial distribution. Both small S. entomon and M. affinis avoided large S. entomon. The avoidance behaviour of M. fffinis contributed about 10 times more to the high proportion in the refuge than numerical effects of predation. Due to the low mortality of small S. entomon the avoidance behaviour of this species was even more important for the spatial distribution. The combined effect of avoidance behaviour and predation in both species was aggregation, producting a positive correlation between the species in density. M. affinis showed two types of avoidance behaviour. In the activity experiments they reduced activity by 36% and buried themselves in the sediment. In the refuge experiments we also observed avoidance behaviour with the emigration rate from the predator compartment being twice the immigration rate. The refuge did not lower predation mortality in M. affinis, probably due to the small scale of the experimental units in relation to the mobility of the species. Predation mortality in small S. entomon was higher in absence of a refuge and especially high in absence of M. affinis.