Surface patterns of zooplankton spatial variability detected by high frequency sampling in the NW Mediterranean. Role of density fronts

High frequency sampling was performed in daylight hours along a 35 km transect in the Ligurian Sea to investigate the upper layer zooplankton distribution during the spring phytoplankton bloom. The results show detailed spatial structure and biomass of key zooplankton functional groups, copepods, salps and krill larvae, within the different water masses characterizing this region. Although observed values of total copepod biomass distribution were rather constant along the transect, species-specific patterns were observed in the copepod spatial distribution. The larger species Calanus helgolandicus, as well as Centropages typicus, Oithona spp., and Oncaea spp., were associated with the frontal zone. However, Acartia spp. had a scattered distribution, and Clausocalanus/Paracalanus did not have a clear pattern. In addition, krill larvae were concentrated in the frontal area and salps had a scattered pattern. The cross-shore zooplankton distribution appeared strongly influenced by both the Northern Ligurian current governing inshore waters, which acts as a major flushing forcing, and the Ligurian front, which governs offshore waters and may act as retention area for zooplankton.     

A numerical investigation of the impact of turbulence on the feeding rates of Oithona davisae

Individual based numerical simulations of the copepod, Oithona davisae, feeding on motile prey, Oxyrrhis marina, under variable turbulent conditions are performed. These simulations correspond to laboratory observations conducted by Saiz et al. [Saiz, E., Calbet, A., and Broglio, E., 2003. Effects of small-scale turbulence on copepods: the case of Oithona Davisae. Limnol. Oceanogr., 48:1304–1311.].The flow field in the simulation is reconstructed by a kinematic simulation whose characteristic scales are derived from the grid mesh and the dissipation rates of the laboratory experiments. The kinematic simulation provides a simplified model, which while not fully realistic, captures the basic relevant feature of turbulence. A hop and sink swimming behaviour is prescribed for O. davisae, while O. marina moves along helical paths with random changes of directions.Three possible effects are tested: the existence of a time threshold in the duration of the contacts between predator and prey, a progressive reduction of the perceptive distance with increasing turbulence level and an abrupt reduction in feeding of O. davisae when the flow speed, in relation to the copepod position, is higher than a prescribed threshold. This last approach introduces an intermittency in the feeding which depends on the variations of velocity both in space and time within the numerical box.The introduction of the time threshold causes a dome-shaped relationship between the simulated enhancement factor and the dissipation rate, while with the other two effects, a monotonic decrease in the enhancement factor is observed, with values reasonably close to the ones observed in the laboratory experiment. In all the cases, the use of realistic values of biological parameters (e.g. swimming behaviour) reproduces response curves in the range of the observations.     

Seasonal carbon distribution of copepods in the eastern Weddell Sea, Antarctica

Copepods were sampled by a multiple opening-closing net in the eastern Weddell Sea during various seasons (late winter/early spring, summer, autumn). Total copepod biomass integrated over the upper 1000 m varied seasonally between 1.7 mg C m⁻³ in late winter/early spring and 3.7 mg C m⁻³ in autumn. After the dark season the copepods were rather evenly distributed vertically and highest biomass levels were found in the mid-water layers between about 200 m and 500 m. By contrast, especially in summer but also in autumn copepod biomass concentrated in the uppermost water layer. A total of 64 calanoid species were identified in the upper 1000 m with maximum species numbers in the deepest layer. The large calanoids Calanus propinquus, Calanoides acutus, Metridia gerlachei, Euchaeta antarctica and the small calanoid Microcalanus pygmaeus prevailed and accounted for 60–70% of total copepod biomass, while the small poecilostomatoid Oncaea and the cyclopoid Oithona species comprised about 20%. Hence, the distribution pattern of the entire copepod biomass is strongly influenced by the life cycles of a few dominant species.     

Annual cycle of distribution of three ice-associated copepods along the coast near Dumont d'Urville, Terre Adélie (Antarctica)

In polar regions sea ice is a site of enhanced primary production during winter and provides important habitat for small grazers, such as copepods. We sampled zooplankton from the sea ice and water column throughout 2005, near Dumont d'Urville station (Terre Adélie, Antarctica). Three species of ice-associated copepods were found: two calanoid copepods Paralabidocera antarctica and Stephos longipes and the harpacticoid copepod Drescheriella glacialis. P. antarctica was the most abundant of the three and was closely associated with the sea ice during most of the year. This species had a one year life cycle with a probable over-wintering period in the sea ice as nauplii and a short copepodite phase in spring. Reproduction and spawning occurred in early summer. A comparison with two other populations (near Syowa and Davis stations) along the east coast of Antarctica showed that there was a temporal shift in the life cycles of the three populations, which was linked to variability in sea ice conditions. D. glacialis was the second most abundant copepod and was more common during the winter than during summer, indicating its preference for the sea ice habitat. In autumn, the presence of D. glacialis in the deeper part of the water column suggested that this species colonised the sea ice from the benthos. S. longipes was found only in the water column near Dumont d'Urville and was not very abundant. In Terre Adélie particular environmental conditions, such as the absence of a permanent sea ice zone throughout the year, a longer time of open water, strong katabatic winds and the presence of polynyas, have influenced both the abundance and distribution of the three common ice-associated copepods.     

Colonization of a new man-made river (Marchfeldcanal, Lower Austria) by benthic copepods

The role of copepods in the colonization and emigration processes of benthos and drift of an irrigation canal were studied. During the 3 first years of existence of the canal, fifteen species of copepods were recorded (1 diaptomid, 11 cyclopoids, 3 harpacticoids). Copepod succession in the canal was initially represented by lentic species, then by predatory species like Macrocyclops albidus that developed with the proliferation of filamentous algae and epibionts during the first two years. Primary consumers appeared from the beginning of the flowing conditions but started to dominate only from the third year. Smaller species associated with the bed sediments like the harpacticoid species Nitocra hibernica, were late colonizers. The latter as well as Paracyclops fimbriatus and Eucyclops serrulatus were the dominant copepod species in the canal. Total copepod density in the benthos ranged from 15,800 to 68,000 ind./m², with a mean of 7.6% of the total zoobenthos. Copepods constituted up to 25% of the total faunal drift density, ranging from 50 to 140 ind./m³. Their highest abundance was observed around midnight. A three-fold increase of the discharge temporarily affected the drift density, but did not change the diel rhythm of the drift. Cyclopoid copepods namely P. fimbriatus (54.77%), copepodid cyclopoids (20.76%) and E. serrulatus (16.11%) formed the main part of the drift samples. The harpacticoid Nitocra hibernica was a minor participant in the drift processes.     

Harpacticoid and cyclopoid fauna of groundwater and springs in southern Finland

The groundwater fauna, of which scarcely anything has previously been studied in formerly glaciated areas, was richest close to the water table and diminished markedly at greater depths. However, few individuals were generally recorded per 1 m³ of water. Bryocamptus minutus was the most abundant species of copepod at an esker site, the other species recorded beingAttheyella crassa, Bryocamptus pygmaeus, Moraria brevipes and Parastenocaris phyllura, and some species from the genus Diacyclops. At a shore bank infiltration site M. brevipes was the most numerous species. However, one esker site examined did not have any animals. Springs, which were grouped into those in a natural state and those under anthropogenic influence, had some species in common with esker groundwater, but also Paracyclops fimbriatus, Acanthocyclops robustus, Acanthocyclops vernalis, Diacyclops bicuspidatus, Megacyclops viridis, and the harpacticoid species Bryocamptus cuspidatus, Bryocamptus echinatus and Canthocamptus staphylinus. Of the spring copepods, P. fimbriatus appeared to withstand the influence of road de-icing with NaCl, but numbers of Moraria brevipes were reduced under this influence. Bryocamptus echinatus was more numerous in springs having a high oxygen content and high pH. The species found in esker groundwaters and springs were in part the same evidently cold-stenotherm species which inhabit the profundal zone of oligotrophic lakes.     

Copepod diversity in the water masses of the southern East China Sea north of Taiwan

The warm oceanic current Kuroshio and the continental shelf water of the East China Sea meet in the western North Pacific, north of Taiwan and form an upwelling when they converge. The intrusion of the Kuroshio westward over the East China Sea shelf thus results in complicated exchanges of waters between these two water masses. We studied the copepods in the plankton collection taken from an east–west transect crossing these waters in April 1995 when the intrusion of the Kuroshio over the East China Sea shelf was beginning to retreat. The taxonomy of copepods was carefully treated and erroneous species records reported in the literature were guarded against. We evaluated the copepod diversity, the association of copepod species, and the association of stations in these water masses.     

Egg production of the copepod Acartia hongi in Kyeonggi Bay, Korea

Many studies of copepod egg production have shown that food availability and temperature are major factors that influence copepod growth. However, coastal environments are complicated ecosystem and the relationships between growth of copepods and influencing factors are not always clear in nature. We conducted a study along an inner–middle–outer bay transect where variations in environmental parameters would be expected to affect the biomass and egg production rate of A. hongi from February 2001 to December 2001. In this study, we investigated the abundance and biomass with developmental stages and egg production rates of A. hongi in relation to various environmental factors. The copepod A. hongi occurred continuously throughout the year, with a peak abundance in May. In general, the variation in egg production rates showed a similar tendency with the variations in chlorophyll-a throughout the study period. This suggests that phytoplankton biomass is an important factor that affects the egg production of A. hongi. In addition, during the warm season, the egg production of A. hongi was also influenced by the ciliates abundance in the middle and outer bay. Consequently, the egg production of A. hongi is generally affected by food availability in Kyeonggi Bay.     

Symbolic dynamics and entropies of copepod behaviour under non-turbulent and turbulent conditions

We consider here behavioural activity of copepods as a succession of symbols associated with swimming states: slow swimming, fast swimming, break and grooming. We characterise these symbolic sequences using tools from symbolic dynamics: probability density function of the residence times in each state; transition probability at each time step; Shannon entropy and dynamic entropy. This approach is applied to the swimming behaviour of Centropages hamatus which we have analyzed as an example of application, in order to stress the differences associated with turbulent and non-turbulent conditions. We characterise in this theoretical framework the behavioural changes exhibited by the copepod under turbulent conditions.     

Variability of mesozooplankton spatial distribution in the North Aegean Sea, as influenced by the Black Sea waters outflow

The North Aegean Sea constitutes an important region of the Mediterranean Sea since in its eastern part the mesotrophic, low salinity and relatively cold water from the Black Sea (outflowing from the Dardanelles strait) meets the oligotrophic, warm and very saline water of Levantine origin, thus forming a thermohaline front. Mesozooplankton samples were collected at discrete layers according to the hydrology of the upper 100 m, during May 1997 and September 1998. In May highest biomass and abundance values (up to 66.82 mg m^− 3 and 14,157 ind m^− 3) were detected in the 10–20 m layer (within the halocline) of the stations positioned close to the Dardanelles strait. The front moved slightly southwards in September, characterized by high biomass and abundance values within the halocline layer. The areas moderately or non influenced by Black Sea water revealed lower standing stock values than the frontal area in both cruises and maxima were detected in the uppermost low salinity layer. Samples collected at the stations and/or layers more influenced by Black Sea water were distinguished from those collected at layers and/or stations more affected by Levantine waters in both periods. In May the former samples were characterized by the copepods Acartia clausi, Centropages typicus, Paracalanus parvus. The abundance of the above species decreased gradually with increasing salinity, in the horizontal and/or in the vertical dimension, with a parallel increase of the copepods Oithona plumifera, Oithona copepodites, Oncaea media, Ctenocalanus vanus, Farranula rostrata. During September the frontal area as well as that covered by the modified Black Sea water, were highly dominated by the cladoceran Penilia avirostris and doliolids. For both seasons, MDS plots, issued from the combination of mesozooplankton and water-type data, revealed the gradual differentiation of zooplankton composition from the frontal area towards the area covered by Levantine water, following the spreading and mixing of the Black sea water. The observed temporal and spatial variability in the distribution pattern of mesozooplankton standing stock and species composition seems to depend considerably on the variability of circulation and frontal flows.     

Intermittent turbulence and copepod dynamics: Increase in encounter rates through preferential concentration

Turbulence has a strong influence on plankton contact rate, which is a crucial parameter for plankton ecology. In the field of particle-turbulence interactions, it is now well known that fully developed turbulence does not always homogenise particle distributions, but instead creates, in some well-defined conditions, preferential concentrations of heavy particles. Many studies have considered the influence of this type of preferential concentration on particle contact rate. We consider here the possible application of these results to copepods, assuming that some results obtained for heavy particles are still valid for light particles. Using parameter values representative of copepod species in coastal waters and open ocean, we numerically estimate the possible enhancement of copepod contact rates due to the preferential concentration by turbulence. The assessment is done by using data from a laboratory experiment: we find from the trajectory analysis of small neutrally buoyant particles, that the preferential concentration effect increases the contact rate up to 140%. We argue that this effect may be more pronounced for higher Reynolds numbers, and may have important ecological applications.     

The role of chemical signals in copepod reproduction

Because the reproductive fitness of a copepod is influenced by its ability to discriminate between conspecific and heterospecific mates, it is important to understand the mechanisms involved in these complex mate-recognition systems. Several lines of evidence suggest that chemical communication plays a role in mate location, recognition and reproduction in copepods. However, the relative importance of chemosensory cues, as well as other factors, including hydromechanical and morphological cues, remains unclear. Some studies show that mating ‘mistakes' occasionally occur in nature, which for species that mate only once, would have considerable effects on fitness. By reducing the likelihood of heterospecific matings, species-specific mate location and recognition chemicals have high adaptive value. The relative importance and molecular characteristics of these chemicals, whether diffusible or contact signals, may explain differences in premating and copulatory behavior observed among different copepod taxa. Current research suggests that both forms of chemical communication significantly influence the reproductive success of copepods. In this review, we summarize examples from the literature that may serve as a means to critically analyze postulated ideas and offer new questions for future study.     

Analysis of the parasitic copepod species richness among Mediterranean fish

The Mediterranean ichthyofauna is composed of 652 species belonging to 405 genera and 117 families. Among these, 182 were studied for their parasitic copepods. The analysis of all the works conducted on these crustacea yielded 226 species distributed in 88 genera and 20 families. For each fish species we have established a file providing the species name of the fish, its family, its geographical distribution within the Mediterranean and some of its bio-ecological characteristics. Within each file, all the parasitic copepod species reported on each host species were listed. This allowed to know the species richness (SR) of these hosts. We thus produced 182 files within which 226 copepod species are distributed. A program was created under the Hypercard software, in order to analyse our data. Two parameters were studied. The first one is the mean species richness (MSR), which corresponds to the mean of the different SR found on the different host species. The second is the parasite–host ratio (P/H), which is the ratio of the number of copepod species by the number of host species. These parameters are calculated by our program for all the 182 species of Mediterranean fishes retained in our investigation, on the first hand, and, on the second hand, for one particular group of fish species. We used the following variables to investigate their correlations with copepod species richness: taxonomy—fish families, genera and species; biometry—maximal size of the adult fish; eco-ethology—mode of life (benthic, pelagic or nectonic), displacements (sedentary, migratory with environmental change, or migratory without environmental change), behaviour (solitary or gregarious). Other variables (colour, food, reproduction, abundance, distribution area) were also analysed but did not reveal any clear correlation. Providing that our study does not rely on quantitative (prevalence, intensity) but qualitative basis our aim was only to reveal some tendencies. These tendencies are as follows: (1) In many cases, parasite and host phylogeny seem to play an important role. There are fish families with copepods and families with few species of these parasites. The phyletic constraints could be due to the morphological characteristics of the habitat (e.g. structure of the gills) or biological/ecological characteristics that we were unable to identify. (2) It appears that the presence in a same environment of related fish species (e.g. several species of the same genus, or numerous genera of the same family) is correlated with high parasite richness. A likely explanation is that such situations favours alternated processes of lateral transfers and speciation. (3) Some eco-ethological criteria seem to favour the establishment of a large parasite species richness. It should be noted for instance that Mediterranean fishes the most often infected with copepods are generally nectonic or pelagic, migratory, and gregarious species.     

Diel changes in vertical overlap between Cyclops strenuus (Copepoda; Cyclopoida) and its prey in oligotrophic Lake Toya, Hokkaido, Japan

The vertical distribution of Cyclops strenuus and its prey, and the gut contents of C. strenuus were investigated at 3 h intervals over 24 h periods in the pelagic area of oligotrophic Lake Toya in May, August and October 1992. C. strenuus showed slight diel vertical migration (DVM) in May, but did not migrate and was always distributed below 15 m in August and October, the thermally stratified period. This suggests that the DVM is strongly influenced by the development of the thermocline, which the copepod may not be able to cross. Gut content analyses revealed that in Lake Toya it is omnivorous, and that the dominant prey are cladocerans and Filinia longiseta. The number of prey remains in the gut varied with time, and the diel changes did not follow changes in prey density that take the vertical overlap between predator and prey into account. We argue that very low prey density associated with the oligotrophic nature of the lake would mask the relationship between prey density and the number of prey in the gut of the predator.     

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological activities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya and due to low ingestion in the remaining regions.     

Mesozooplankton species distribution in the NW and N Iberian shelf during spring 2004: Relationship with frontal structures

We have analysed the mesozooplankton community structure in the southern Bay of Biscay shelf and its relationship with the hydrographic conditions during spring 2004. According to thermohaline characteristics, we observed two frontal zones of distinct origin along the shelf (around 7° and 3°W), that allowed us to differentiate three different hydrographic domains. The westernmost part of the shelf (WC), defined by the presence of relatively warm and salty water related to the presence of the Iberian Poleward Current (IPC), the easternmost region (EC), characterized by colder and fresher water and subject to the influence of freshwater inputs from the Adour river in the French coast, and a region in the Central Cantabrian Sea (CC), where thermohaline characteristics were intermediate between these two extremes. The mixing layer depth (MLD) regime in these areas was also different: the WC region was characterized by a mixed water column, whereas in the EC region the river discharges produces stratification of the upper meters of the water column (< 10 m); in the CC region, we found a distinct vertical mixing regime that separated coastal (stratification) from shelf (mixed water column) stations, giving rise to a notorious across-shelf front. We found a good match between the aforesaid hydrographic regions and the distribution of mesozooplankton species composition and community assemblages: the Mantel correlation between physical variables and mesozooplankton distribution was highly significant (n = 63, r = 0.70, α < 0.001). In the WC region, the community was dominated by Paracalanus parvus, Oithona helgolandica, Acartia clausi and Clausocalanus pergens, while in the EC region the most dominant species were Noctiluca scintillans, Oncaea media and Temora longicornis. The CC region showed similar composition of copepods than the WC region, but larvaceans (Oikopleura spp. and Fritillaria spp.) were more abundant in the CC region than in the WC region. Within each zone, the relative abundances of the dominant species differed between coastal and shelf locations.     

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus, were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m⁻² and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis, inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus, where grazing is negligible, at least during early summer.     

Effects of small-scale turbulence on the growth of two diatoms of different size in a phosphorus-limited medium

The effect of turbulence on the nutrient flux towards osmotrophic cells is predicted to be size dependent. This should translate into growth. We experimentally followed and modelled the growth of two marine diatoms of different size (Thalassiosira pseudonana, 6 μm in diameter and Coscinodiscus sp., ca. 109 μm in diameter) under still water and turbulent conditions, using a shaker table. Experiments were done with phosphorus-limited cultures and lasted for ca. 5 days. Turbulence enhanced the growth of Coscinodiscus sp. in agreement with theory but not the growth of T. pseudonana, which was actually slightly lower under turbulence. At the end of the experiments there were about 1.7 times as many Coscinodiscus sp. cells in the turbulent treatment than in the still treatment, while for T. pseudonana almost the same cell concentration was found in both conditions. In addition, the Coscinodiscus sp. cells growing under still conditions presented a higher specific alkaline phosphatase activity than those growing in turbulence which indicates a higher need for phosphorus in the still cultures. A simple dynamic model, based on Michaelis–Menten nutrient uptake kinetics, needed nearly no optimisation other than using observed initial conditions of phosphate and cell concentrations. The model showed how an increased nutrient flux towards the cells translates non-linearly into cell growth, most likely by affecting the half-saturation constant (K_M). However, since Coscinodiscus sp. experienced significant mortality and cells partially settled to the bottom of the containers, unequivocal support for the size-dependent effect of turbulence on nutrient uptake will require further experiments and more sophisticated modelling. The mechanisms to connect an increased nutrient flux towards cells with population growth and whether this process is size dependent are important in parameterizing the effects of turbulence on marine plankton in coupled physical–biological models.     

航道清礁对西江鱼类保护区紊流体的影响

河底裸露的礁石增加水体紊动,有助于鱼卵正常形成,大量存在礁石的河段往往是鱼类栖息和产卵的重要场所。礁石也存在碍航的可能,航道清礁一定程度上影响水体紊动强度,改变鱼类栖息环境。为此,在重要鱼类栖息河段进行航道清礁需充分论证。以紊动动能作为评价水体紊动强度指标,开展西江中游珍稀鱼类保护区河段三维紊流数值模拟,计算并统计航道清礁工程前后水体紊动动能及不同强度的紊流水体体积,分析其变化规律及原因。研究结果表明,断面中紊动动能强度与水深成正相关关系;清礁工程导致河段中弱紊动动能的紊流水体体积增加,强紊动动能的紊流体积减小;底层水体紊动动能的变化量高于表层;强紊动动能水体体积减少的变化率大于弱紊动动能水体体积增加的变化率。