Small scale retentive structures and Dinophysis

Despite its rarity, Dinophysis acuminata is in terms of economic impact, the first toxic algal species along the coasts of Western Europe. It is observed at low levels (< 20 cell l^− 1) all the year round but toxic events occur mainly in late spring and summer. D. acuminata ecophysiology is largely unknown due to the inability to culture it. Therefore, standard biomass models based on inorganic nutrition are largely inadequate. Presently, any progress in describing the conditions of population growth of this species will be a step forward to prediction of harmful events at the coast. This species has been observed at increased, albeit low cell densities in retentive eddies located in pycnocline layers. A concentration build-up of one species results from the balance between growth and loss processes, one of the latter being dispersal. The scales of interest for a D. acuminata population are of the order of 10 nautical miles on the horizontal and duration of 10 days, for a reported achievable growth rate of 0.6 day^− 1. A three dimensional (3D) hydrodynamical model of the Bay of Biscay has been elaborated to reproduce hydrological structures over the last decade. We attempt here to relate the existence of retentive structures revealed from simulations under realistic forcing conditions and the toxic coastal events recorded in the 10-year time series of the French plankton monitoring network database. The eddies in the coastal area appear to be directly related with the Dinophysis coastal events and they may be a potential effective tool to predict those.     

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.     

Haplostoma dudleyae sp. nov. (Cyclopoida: Ascidicolidae), parasitic in Eudistoma olivaceum from the Indian River in southern Florida

Haplostoma dudleyae sp. nov. is described and illustrated on the basis of females found in a compound ascidian (Eudistoma olivaceum) collected from the Indian River in southern Florida, USA. Each parasitized zooid has a single female copepod. The egg sacs of the female are paired and, when laid, are almost U-shaped and folded against the dorsal side of the body. Haplostoma dudleyae and two other species (H. canui Chatton and Harant, 1924, and H. humesi Ooishi, 1995) constitute a subgroup within the genus Haplostoma.     

New species of the genus Xanthocalanus (Copepoda, Calanoida, Phaennidae) from the Laptev Sea

Sampling of the near-bottom calanoid copepods was performed on board of the research vessel Polarstern in 1993 in the Laptev Sea (Siberian Arctic). Three new species of Xanthocalanus were identified as such, and are described and figured here: X. spinodenticulatus sp. nov., X. laptevorum sp. nov. and X. polarsternae sp. nov.     

The gonad morphology and maturation in Arctic Calanus species

Gonad morphology and maturation of the dominant Arctic copepod species Calanus glacialis and C. hyperboreus were studied using a combination of observations on whole animals (“whole-body analysis”) and histological sections to describe basic reproductive processes. The comparison with C. finmarchicus shows that the internal organisation of the gonads is the same in the Calanus species: from the dorsally laying ovary, two anterior diverticula extent into the head region and two posterior diverticula extend into the abdomen. In the diverticula, the oocytes increase in size and development stage in a dorso–ventral direction. There were no physiological differences between the oocytes in the diverticula and oviducts in any of the three species. During gonad development, the morphological modifications of the gonads mainly refer to an increase in number, developmental stage and size of the oocytes both in diverticula and oviducts. During spawning the most ventral layer of oocytes is released, and then the adjacent dorsal layer begins final maturation. Thus, the final maturation processes occur simultaneously as it is essential to produce clutches.     

Observations on colony formation by the cosmopolitan phytoplankton genus Phaeocystis

Few marine phytoplankton have heteromorphic life cycles and also often dominate the ecosystems in which they occur. The class Prymnesiophyceae contains a notable exception: the genus Phaeocystis includes three species that form gelatinous colonies but also occur within their ranges as solitary cells. Phaeocystis antarctica and P. pouchetii are exclusively high latitude taxa, and are notable for regionally tremendous blooms of the colony stage. P. globosa occurs circumglobally, yet its colony blooms primarily are confined to colder waters within its range. Three additional species are warm water forms that have been reported only as solitary cells or loose aggregations that bear little resemblance to the organized colonies of the other taxa. Interpretation of existing data indicates that resource availability (light, temperature and nutrients) by itself is not sufficient to explain this distinction between cold-water colony-forming taxa and warm water solitary cell taxa, nor why colony development in P. globosa is essentially a spatially restricted phenomenon within a much broader geographic range. Colony development by P. globosa in situ has been observed at temperatures ≥20 °C, but only rarely and generally under conditions of seasonally or anthropogenically elevated nutrient supply. Data presented here demonstrate colony development at 20–22 °C in natural plankton communities from oligotrophic waters that were pre-screened through 63 μm mesh (i.e. lacking mesozooplankton and large microzooplankton), but not in unscreened communities containing microzooplankton and >63 μm zooplankton. Reduction of colony proliferation at higher temperatures by mesozooplankton grazing remains as an intriguing possibility that is consistent with available evidence to help explain differences in latitudinal extent of in situ colony development. These data are interpreted within a theoretical framework regarding the potential advantages and disadvantages of the two life cycle stages.     

The female antennules of Paracyclops (Cyclopoida: Cyclopidae): their significance for systematics

The setation patterns of female antennules of some Paracyclops species are analysed with reference to the hypothetical 28-segmented antennule of ancestral copepod proposed by Huys and Boxshall in 1991. Comparison between Paracyclops species has revealed a setation pattern common to all species thus far examined. Several modifications of the female antennules have been identified as apomorphic states and can be used in phylogenetic studies. P. waiariki has 12-segmented antennules and exhibits more plesiomorphic characters than the reduced 8-segmented antennules of other species, such as P. fimbriatus. The results confirm that simple segment numbers are unusable as characters in cladistic analysis. Both P. affinis and P. yeatmani, for example, have 11-segmented antennules but analysis of segmental homologies has revealed that the 11-segmented state is convergent in these two species.     

Diel vertical migration and feeding in adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani during a spring bloom in Dabob Bay, a fjord in Washington USA

Diel vertical migration and feeding on phytoplankton by adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani were simultaneously measured near the end of a phytoplankton bloom. Almost the entire Calanus population migrated out of the deep layer (108–50 m) at night but only about 30% came to the surface (25–0 m). Feeding occurred only at night and was equally high in the surface and intermediate layers, in spite of much higher food concentrations in the surface. Like Calanus, the entire Metridia population was found in the deep layer during the day but unlike Calanus, 20–50% remained in the deep layer at night and most migratory Metridia were collected from the surface layer. Metridia feeding at night was highest in the surface layer but significant feeding also occurred in both the intermediate and deep layers. Migratory behavior of Pseudocalanus was weak, with the proportion of the population in the surface layer increasing from slightly <10% during the day to approx 30% at night. Feeding occurred in both surface and intermediate layers throughout the 24 h but was greater in both layers at night. The different migratory patterns are discussed in the context of our current understanding of the contributions of predator avoidance and feeding to diel vertical migration.     

Application of new parameterizations of gas transfer velocity and their impact on regional and global marine CO2 budgets

One of the dominant sources of uncertainty in the calculation of air–sea flux of carbon dioxide on a global scale originates from the various parameterizations of the gas transfer velocity, k, that are in use. Whilst it is undisputed that most of these parameterizations have shortcomings and neglect processes which influence air–sea gas exchange and do not scale with wind speed alone, there is no general agreement about their relative accuracy.The most widely used parameterizations are based on non-linear functions of wind speed and, to a lesser extent, on sea surface temperature and salinity. Processes such as surface film damping and whitecapping are known to have an effect on air–sea exchange. More recently published parameterizations use friction velocity, sea surface roughness, and significant wave height. These new parameters can account to some extent for processes such as film damping and whitecapping and could potentially explain the spread of wind-speed based transfer velocities published in the literature.We combine some of the principles of two recently published k parameterizations [Glover, D.M., Frew, N.M., McCue, S.J. and Bock, E.J., 2002. A multiyear time series of global gas transfer velocity from the TOPEX dual frequency, normalized radar backscatter algorithm. In: Donelan, M.A., Drennan, W.M., Saltzman, E.S., and Wanninkhof, R. (Eds.), Gas Transfer at Water Surfaces, Geophys. Monograph 127. AGU,Washington, DC, 325–331; Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] to calculate k as the sum of a linear function of total mean square slope of the sea surface and a wave breaking parameter. This separates contributions from direct and bubble-mediated gas transfer as suggested by Woolf [Woolf, D.K., 2005. Parameterization of gas transfer velocities and sea-state dependent wave breaking. Tellus, 57B: 87–94] and allows us to quantify contributions from these two processes independently.We then apply our parameterization to a monthly TOPEX altimeter gridded 1.5° × 1.5° data set and compare our results to transfer velocities calculated using the popular wind-based k parameterizations by Wanninkhof [Wanninkhof, R., 1992. Relationship between wind speed and gas exchange over the ocean. J. Geophys. Res., 97: 7373–7382.] and Wanninkhof and McGillis [Wanninkhof, R. and McGillis, W., 1999. A cubic relationship between air−sea CO2 exchange and wind speed. Geophys. Res. Lett., 26(13): 1889–1892]. We show that despite good agreement of the globally averaged transfer velocities, global and regional fluxes differ by up to 100%. These discrepancies are a result of different spatio-temporal distributions of the processes involved in the parameterizations of k, indicating the importance of wave field parameters and a need for further validation.     

Identification and quantification of plankton bloom events in the Baltic Sea by continuous pCO2 and chlorophyll a measurements on a cargo ship

Continuous measurements of the surface water CO₂ partial pressure (pCO₂) and the chlorophyll a fluorescence were performed in the Baltic Sea using a fully automated measurement system deployed on a cargo ship. The ship commuted regularly at two day intervals between the Mecklenburg Bight (Luebeck) and the Gulf of Finland (Helsinki). The pCO₂ data collected during June 2003 and September 2004 were used to identify biological production events such as the spring bloom and the midsummer cyanobacteria bloom in five different sub-regions. To quantify the net biomass production, the decrease of the total CO₂, ^NC_T (normalized to a uniform alkalinity), during the production periods was calculated using the pCO₂, temperature and salinity records and the mean alkalinity. Taking into account the CO₂ air/sea exchange and the formation of dissolved organic carbon, a simple mass balance yielded the net production of particulate organic carbon which represents the total biomass. The chlorophyll a concentrations obtained from the fluorescence data showed peaks that in most cases coincided with the production maxima and thus supported the interpretation of the pCO₂ data. The production during both the spring bloom (2004) and the midsummer nitrogen fixation period (2003) increased by a factor of about three from the southwest to the northeast. For the spring bloom our estimates were significantly higher than those based on the winter nutrient supply and Redfield C / N and C / P ratios. This indicated the existence of additional nutrient sources such as dissolved organic nitrogen, early nitrogen fixation and preferential P mineralization. Midsummer ^NC_T minima were observed only in 2003 and used to quantify the nitrogen fixation activity and to characterize its interannual variability.     

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.     

Limitation of egg production in Calanus finmarchicus in the field: A stoichiometric analysis

The egg production of marine copepods correlates with a range of variables, including the availability of organic carbon (C), nitrogen (N) and the polyunsaturated fatty acids (PUFAs) 20:5(n−3) (EPA) and 22:6(n−3) (DHA). However, an understanding of which substrates limit egg production in the natural environment has yet to be reached. The quantities of C, N, EPA and DHA ingested, derived from parental biomass, and invested in eggs by female Calanus finmarchicus during a 5-day incubation experiment were examined using stoichiometric theory to determine which substrate was limiting. The majority of each substrate was derived from parental biomass, and therefore the existing stoichiometric theory is developed to include this route of supply. The females were essentially devoid of lipid reserves, as evidenced by the lack of the storage fatty acids 20:1(n−9) and 22:1(n−11), and carbon limitation was predicted under most of the scenarios examined. Nitrogen limitation was only apparent when carbon and nitrogen utilisation efficiencies were assumed to be high (0.5) and low (0.4) respectively. PUFAs were assumed to be utilised with high efficiency (0.9), and were never predicted to limit production. This work highlights the need for a more detailed understanding of the maintenance requirements that marine copepods have for C, N, EPA, and DHA and hence the efficiencies with these substrates can be utilised for growth.     

Appendicularian ecophysiology I: Food concentration dependent clearance rate, assimilation efficiency, growth and reproduction of Oikopleura dioica

Three aspects of the appendicularian O. dioica's ecophysiology were measured here: 1) morphological parameters over a wide range of appendicularian sizes, including mature animals in order to document the morphological characteristics inducing reproduction; 2) clearance rate and assimilation efficiency using feeding incubations with different algal concentrations and 3) the effect of food concentration on growth, mortality and reproduction.The relationship between the body carbon weight and the clearance rate follows a power function, with an exponent of 0.91 (± 0.07). The rate of particles retention increases with the food concentration following a Michaelis–Menten relationship (half-saturation constant = 151 ± 22 µg C l^− 1, maximum clearance rate = 12 ± 1 µg C µg C^− 1 d^− 1). The carbon assimilation efficiency decreases with the increasing food concentration. As a result, appendicularian growth which is limited in concentrations lower than 50 µg C l^− 1 is saturated above 100 µg C l^− 1.In immature animals the gonad represents less than 30% of the body volume whereas in mature individuals, its volume varies between 50% and 87% (mean 63%) suggesting that gonad/total volume ratio can be used as indicator of the maturation stages. The gonad weight in mature animals represents 70.3 (± 4.6)% of the total body carbon weight. Two major maturity stages can explain the changes in energy allocation: i) the somatic growth, when less energy is invested in gonad growth when compared to the rest of the body and ii) the maturation phase where most of the assimilated matter is invested in gonad maturation. This process is rapid, lasting only few hours. For this reason we measured completely mature organisms that are generally not measured during the experimental work with appendicularians. In food-limited conditions, the gonad maturation process starts with smaller individuals and ends with smaller reproductive animals having the same gonad to total volume ratio than in unlimited food conditions. The results obtained in this study were used to model the life cycle of O. dioica (see Lombard, F., Sciandra, A. and Gorsky, G., 2009-this volume. Appendicularian ecophysiology. II. Modeling nutrition, metabolism, growth and reproduction of the appendicularian Oikopleura dioica.).     

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.     

Additions to Mexican freshwater copepods with the description of the female Leptodiaptomus mexicanus (Marsh)

A survey on 10 temporary and permanent water-bodies, located at more than 2000 m above the sea level in the central part of Mexico resulted in the presence of 17 cyclopoid and five calanoid copepods. From the total taxa found, seven are new to the country: Leptodiaptomus assiniboiaensis (Anderson), Microcyclops dubitabilis Kiefer, Ectocyclops rubescens Brady, Eucyplops cf. bondi Kiefer, Eucyclops cf. prionophorus Kiefer, Eucyclops pseudoensifer Dussart, and Eucyclops cf. solitarius. The female of Leptodiaptomus mexicanus, a species known only from a single male found near Mexico City in 1929, is here described for the first time. Copepod fauna for each of the systems was highly diverse, with up to 12 species co-occurring in temporary ponds. This study revealed a mixture of north and south American taxa in the area.     

Pigments, size and distribution of Synechococcus spp. in the Black Sea

Pigments, size and distribution of Phycoerythrin-containing unicellular cyanobacteria Synechococcus spp. within the euphotic zone were studied for the first time in April–May 1994 in the western and southwestern Black Sea by epifluorescence microscopy and flow-cytometry. Synechococcus was present in varying quantities at every station and depth studied. Surface spatial distribution of Synechococcus revealed that cells were much more abundant in offshore waters than near coastal regions under the direct influence of the Danube river. Minimum and maximum cell concentrations ranged between 9×10² and 1.45×10⁵ cells/ml at the surface, between 2×10³ and 1.23×10⁵ cells/ml at the chlorophyll sub-maximum layer, and between 1.3×10² and 3.5×10² at the nitrite maximum layer. Cells at the chlorophyll sub-maximum layer (based on in-situ fluorometer readings) fluoresce brighter and longer than the ones at the surface and lower depths. Spectral properties of chromophore pigment types of total 64 clonal isolates from different depths down to the lower layer of the euphotic zone (60 m) in the southern Black Sea coast revealed that all have type 2 phycoerythrobilin in common, lacking in phycourobilin. In vivo fluorescence emission maxima for the phycoerythrobilin were about the same (578 nm) for all isolates. All isolates examined showed in vivo absorption maxima at between 435 and 442 nm and at about 681 nm due to chlorophyll-a. Based on the flow cytometer mean forward light scatter data for size distribution, it could be concluded that cells at the surface mixed layer (0–10 m) were larger in cell size than the cells at lower depths (20–60 m).     

The trophic role of the tunicate Salpa thompsoni in the Antarctic marine ecosystem

During a repeat grid survey and drogue study carried out in austral summer 1994/95, the abundance and feeding activity of salps were estimated in the Lazarev Sea region from net tows and in situ measurements of gut fluorescence. Throughout the survey area, Salpa thompsoni accounted for >95% of the total salp stock while Ihlea racovitzai was consistently represented in very low abundances. Maximum densities of S. thompsoni, with ≈4000 ind. 1000 m⁻³, were recorded in the Marginal Ice Zone (MIZ) in December when chlorophyll-a concentrations were well below 1 mg m⁻³. A dramatic decrease in salp stock was observed at the beginning of January, when S. thompsoni virtually disappeared from the most productive area of the MIZ where chlorophyll-a concentrations had by then reached bloom levels of 1.5–3 mg (Chl-a) m⁻³. In situ grazing measurements showed that throughout the cruise S. thompsoni exhibited the highest ingestion rates per individual of any of the most abundant components of the grazing pelagic community, with maxima of ≈160 μg (pigm) ind. ⁻¹ d⁻¹. These feeding rates are 3 to 5 times higher than those previously obtained using in vitro incubations. The total daily consumption of the population of S. thompsoni varied from 0.3 to 108% of daily primary production. We suggest that competitive removal of food by S. thompsoni, rather than direct predation, is responsible for the low krill abundances generally associated with salp swarms.     

The influence of food quality on the nutritional acclimation of the copepod Acartia clausi

The influence of food quality on the nutritional metabolism of Acartia clausi was studied experimentally using four different diets: (1) diatom cells of Thalassiosira weissflogii, (2) detritus prepared from the same culture, (3) 50 : 50 mix on a protein basis of the two previous diets, and 4) dinoflagellate cells of Prorocentrum micans. For each trophic, ingestion, gut transit time, trypsin activity and K_m (half saturation constant) were measured at limiting and saturating concentration. Assimilation rates were also estimated for both pure diatoms and mixed live–detrital cell diets. Ingestion followed a Holling type 2 response for diets 1 and 4, a linear one for detritus and an intermediate response for diet 3. Gut transit time displayed different adaptive changes with food regime depending on protein concentration. Trypsin activity was lower for detrital food and trypsin K_m significantly decreased with increasing concentration of live diatoms. Assimilation rates were higher for live food than for mixed live–detrital food. Results illustrated that Acartia-type copepods optimize nitrogen or protein uptake. They suggest that besides chemoreception-mediated selectivity, internal controls by digestion and assimilation also regulate ingestion (feed-back).